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HARVARD UNIVERSITY 




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LIBRARY 



OF THE 



Museum of Comparative Zoology 



APR 12 H 



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BULLETIN OF THE UNIVERSITY OF KANSAS 

Vol. XXIII DECEMBER, 1922 No. 18 

Science Bulletin 

Vol. XIV, Nos. 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 
15, 16, 17, 18, 19, 20 and 21 

(Continuation of Kansas University Quarterly.) 




ENTOMOLOGY NUMBER V 



LAWRENCE, KANSAS 

Published Semimonthly from Januaiy to June and Monthly from July to 
December, inclusive, by the University of Kansas. 



Entered as second-class matter December 29, 1910, at the post oflBce at Lawrence, Kansas, 

under the act of July 16, 1894. 



9-4522 



NOTICE TO EXCHANGES. 

The attention of learned societies and other institutions which 
exchange . s(^ie!ntifi|C publications with the University of Kansas is 
called to the list of publications of this University on the third and 
fourth pages of the cover of this issue. 

Those marked ''Supply exhausted" cannot be furnished at all ; as 
far as the supply permits the remaining numbers will gladly be fur- 
nished to any of our exchanges who may need them to complete their 
files. 

Back numbers of the Kansas University Quarterly, as far as pos- 
sible, will be sent to those of our newer correspondents who are able 
and willing to reciprocate. 



ANNOUNCEMENT. 

The Kansas University Science Bulletin (continuation of the 
Kansas University Quarterly) is issued in parts at irregular inter- 
vals. Each volume contains from 300 to 400 pages of reading-mat- 
ter, with necessary illustrations. Exchanges with other institutions 
and learned societies everywhere are solicited. All exchanges should 
be addressed to the Library of the University of Kansas. 

All communications should be addressed to 

The Kansas University Science Bulletin, 
Library of the University of Kansas, 
Lawrence, Kan. 



EDITORIAL BOARD. 

S. J.. Htjntbb,; Chairman. O. O. Stoland. 

R. C. MootiE, Secretary. W. S. Hunter. 

W. C. Stevens. W. J. Baumgabtner. 

H. C. Teacy. 

Editorial Note. — ^This volume was assembled and published under the edi- 
torial supervision of Dr. H. B. Hungebford. 



APR 12 1924 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



L . n. 



DEVOTED TO 

THE PUBLICATION OF THE RESULTS OF 

RESEARCH BY MEMBERS OF THE 

UNIVERSITY OF KANSAS 



Vol. XIV 

ENTOMOLOGY NUMBER V. 



PUBLISHED BY THE UNIVERSITY, 

LAWRENCE, KANSAS. 

1922. 

9-4522 



CONTENTS OF VOLUME XIV. 



No. PaRo 

1. Historical Account of Department of Entomology 
(continued from Science Bulletin VIII). H. B. 
Hunger ford 9 

2. Withdrawn from publication. 

3. The Membracidae of Kansas (Homoptera). Plates 
I-VII. P. B. Laivso7i 27 

4. The Genus Acf)?op^e?'i4s (Homoptera). Plates VIII- 
XII. P. B. Laivson Ill 

5. The Life History of the Toad Bug (Heteroptera). 
Plates XIII-XIV. H. B. Himgerford 143 

6. A New Subterranean Isopod (Crustacea). Plate 
XV. H. B. Hjingerford 173 

7. Studies in Cicadella hieroghiphica (Homoptera). 
Plates XVI-XX. Lucy M. Hackman 185 

8. Ovipositors of Cicadellidse (Homoptera). Plates 
XXI-XXXIII. P. A. Readio 213 

9. Life History Notes on Two Species of Saldidae (Het- 
eroptera). Plates XXXIV-XXXV. Grace OUve 
Wiley 299 

10. A Problem in the Relation of Temperature to Rate 

of Insect Development. P. A. Glenn 315 

11. Some Biological Notes on Philippine Zoology. F. X. 
Williams 327 

12. Notes on Nesting of Polistea (Hymenoptera). 
Divight Isely 339 

13. Five New Species Belonging to Genus HormoUta 
(Hymenoptera). Plates XXXVI-XXXVII. W. J. 
Phillips and Fred W. Poos 347 

14. The Urinary System of Phlegethontius sexta (Lepi- 
doptera). Plate XXXVIII. George H. Vansell 363 

15. A Brief Resume of Investigations Made in 1913 on 
Trogoderma inclusa (Coleoptera). Plates XXXIX- 
XL. A. H. Beyer 371 

16. The Larva of a Chironomid (Diptera). Plates XLI- 
XLIII. P. W. Claassen 393 

(5) 



6 The University Science Bulletin 

17. Water Insects from a Portion of the Southern Utah 
Desert. R. C. Moore and H. B. Himgerford 407 

18. The Nepidse in America North of Mexico (Heterop- 
tera) . Plates XLIV-LI. H. B. Hungerford 423 

19. A Study of the Relation Between Function and 
Growth in Body Cells. Plates LII. M. T. Burrows 473 

20. Notes on the Biology of Curicta (Heteroptera). 
Plate LIII. Mrs. Grace Olive Wiley 505 

21. Biology and Morphology of Lepyronia quadrangu- 
laris (Say) — Homoptera, Cercopidse. Plates LIV 

to LXII. Kathleen Doering 513 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 1— October, 1922. 

(Whole Series, Vol. XXIV, No. 1.) 

ENTOMOLOGY NUMBER V 



CONTENTS: 

Historical Account of Department of Entomology 
(continued from Science Bulletin VIII). 

H. B. Hungerford. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 1. 



Historical Account of Department of Entomology. 

(Continued from Science Bulletin VIII), 

Brief resume of the work of the Department of Entomology of the University 
of Kansas during the past quarter centuiy. 

BY H. B. HUNGERFORD. 

ENTOMOLOGICAL work has been in progress at the University 
of Kansas since the foundation of the institution in 1866. The 
first faculty consisted of three men, one of whom was Dr. Francis 
Huntington Snow, professor of mathematics and natural science. 

Doctor Snow, while widely interested in birds and flowers, gave 
early evidence of a special fondness for the study of insects. 
Through his efforts, and those of others who have followed him, 
there has been established what is perhaps the greatest general 
collection of insects to be found in connection with any state uni- 
versity in America. 

In 1890 Doctor Snow became chancellor of the University, and 
Dr. Vernon L. Kellogg was appointed to the entomology work, first 
as assistant professor of entomology, and later as associate pro- 
fessor. 

Doctor Kellogg was called to Stanford University in 1894, and for 
the year 1895 the entomological work was in charge of W. A. Snow, 
son of the chancellor. ^ 

In 1896 Prof. S. J. Hunter was appointed assistant professor of 
entomolog}^ and placed in charge of the department, and for the 
past quarter of a century he has directed his energies towiird the 
development of a department that should rank among the strongest 
in the country. During this span of years the department has 
trained many students and grown remarkably in its material equip- 
ment, both through the greatly enriched entomological collections 
and its mechanical facilities for furthering research and advancing 
instruction in entomology. 

(9) 



10 The University Science Bulletin. 

The teaching staff has increased from one to five, and the student 
roll, from a few to more than 250. A total of 2,000 pages, under 
276 titles, have been published by members of the department, 
and twenty-three scientific expeditions of the entomological museum 
have been made during the past twenty-five years. These accom- 
plishments, together with the various economic state problems that 
have arisen from time to time, indicate an active and productive 
period in the history of the department. 

In 1914, Science Bulletin issued its second entomological num- 
ber. This was dedicated to Doctor Snow, and in it may be found 
a chronological account of the activities of the department up to 
and including the year 1913. As a matter of record, additions are 
made below to the various sections as they have occurred since 
that year. 

CHRONOLOGICAL REVIEW OF EVENTS SINCE 1913. 

1913. Mr. George Collett appointed Fellow in Entomology. 

F. X. Williams, who received his M. A. degree in this department in 
1913, resigns to study for the doctorate at Harvard. 

Professor Hunter, Assistant Professor Hungerford, Mr. George Vansell 
and Mr. George Collett conduct a biological survey in Wyoming, 
Utah and Montana with special reference to distribution and biology 
of grasshoppers of Kansas. 

1914. P. W. Claassen becomes assistant state entomologist. 
Raymond Beamer becomes assistant curator of museum. 

Professor Hunter, Assistant Professor Hungerford, Raymond Beamer, 
Will Brown and Fred Poos collect along the Rio Grande in south- 
western Colorado. 

1915. Walter Wellhouse elected fellow in entomology. 

Mr. Raymond Beamer, B. P. Young, Forrest Anderson and Walter Well- 
house make an entomological survey of the counties of southeastern 
Kansas. 

B. P. Young takes charge of the inseetary. 

1916. Forrest Anderson elected fellow in entomology. 
Professor Hunter takes a four months' lea^•e of absence. 

H. B. Hungerford appointed temporary chairman and acting state en- 
tomologist. 

Hungerford, Lawson and Wellhouse go to Welhngton, Kan., to investi- 
gate green-bug outbreak. The Federal Bureau, the Kansas Agricul- 
tural College and the University combine to study the green bug 
and methods of control. Lawson and Wellhouse represent the Uni- 
versity for several weeks in this work. 

Mr. Raymond Beamer, with Grutzmacher, Darby and Gardner, make 
an entomological survey of some southern Kansas counties. 

H. B. Hungerford is granted a year's leave of absence to study for the 
doctorate at Cornell LTniversity. 



Historical Account. 11 

P. W. Claat;!?en takes Mr. HungerfonlV place for the year. 
P. B. Lawson becomes assi-stant instructor. 

1917. H. B. Hungerford inad(> associate professor of entomology. 
P. B. Lawson made instructor in entomology. 

Ruby Hosford takes charge of inscctary. 

Dr. C. P. Alexander becomes assistant curator of entomological museum. 

Miss Eva Batchman appointed assistant instructor. 

William Hoffmann elected fellow in entomology. 

Doctor Ale.xander conducts a party, consisting of Harry Fackler, Wil- 
liam HotTmann and Scott Johnson, on an entomological survey of 
southwestern Kansas. 

Canker-worm work in cities of eastern Kansas conducted by Professor 
Hunter. 

1918. p]ntire University reorganized to handle S. A. T. C. Department of 

entomology teaches three sections of biology. Miss Weaverling takes 
charge of insectary. 

1919. William Hoffmann is apiiointed assistant in charge of insectary. 

P. B. Lawson granted degree of doctor of philosophy' by Kansas LTni- 
versity; major, insect taxonomy; first minor, insect morphology; 
second minor, systematic botany. Title major thesis, "Cicadellidae 
of Kansas." 

Miss Itasca Hilsman elected Fellow in Entomology. 

1920. H. B. Hungerford made Professor of Entomology. 

P. B. Lawson becomes Assistant Professor of Entomology. 
Mr. William Hoffmann transferred from the insectary to assistant cura- 
tor of museum. 
Mr. Philip A. Readio is appointed instructor in entomology. 
Miss Lucy Hackman becomes assistant instructor. 
Miss Itasca Hilsman reappointed fellow in entomology. 

1921. Mr. Lawson becomes associate professor of entomology. 
W. J. Brown becomes student assistant in the museum. 

W. J. Brown, Jean Linsdale and Robert Guentert continue the biological 

survey in northeastern Kansas. 
P. B. Lawson and Raymond Beamer devote some time to the study of 

the destructive outbreak of pea aphis in first-crop alfalfa. 

1922. Miss Kathleen Doering appointed fellow in entomology. 

P. B. Lawson becomes professor of entomology and assistant dean of 
the College of Liberal Arts and Science. 

Miss Kathleen Doering appointed scientific illustrator. 

C. Howard Curran, of Orillia. Canada, appointed research fellow in 
entomology. 

R. H. Beamer reappointed assistant curator of the entomological col- 
lections. 



12 The University Science Bulletin. 

ENTOMOLOGICAL PAPERS FROM UNIVERSITY OF KANSAS. 

(Continued from Science Bulletin VIII.) 
ALEXANDER, C. P. 

No. Date. Title of paper. 

203. 1918 — Records of Japanese Crane Flies (Diptera). Am. Ent. Soc. Amer., 

vol. XI, pp. 443-449. 

204. 1918 — New Species of Tipuline Crane Flies from Eastern Asia (Tipulidse, 

Diptera). Jour. N. Y. Ent. Soc. vol. XXVI, pp. 66-75.- 

205. 1918 — A New Interpretation of the Wing Venation of the Pedicine Crane 

Flies (Tipulidse, Diptera). Ent. News, vol. XXIX, pp. 201-205. 

206. 1918 — New Species of Crane Flies from California (Diptera). Ent. News, 

vol. XXIX, pp. 285-288. 

207. 1918— New Nearctic Crane Flies (Tipulidse, Diptera), Part IV. Can. 

Ent., vol. L, pp. 60-71. 

208. 1918— New Nearctic Crane Flies (Tipulidse, Diptera), Part V. Can. Ent., 

vol. L, pp. 158-165, 242-246. 

209. 1918— New Nearctic Crane Flies (Tipulidse, Diptera), Part VI. Can. 

Ent., vol. L, pp. 381-386, 411-416. 

210. 1919 — The Crane Flies Collected by the Canadian Arctic Expedition, 

1913-"18, Report of the Canadian Arctic E.xpedition, 1913-'18, vol. 
Ill, part C, pp. IC to 30C. 

211. 1919 — Notes on the Genus Dicranoptycha Osten Sacken. Ent. News, vol. 

XXX, pp. 19-22. 

212. 1919 — New or Little-known Crane Flies from Japan. Part I. The En- 

tomological Magazine, Kyoto, Japan, vol. Ill, pp. 122-127. 

213. 1919— The Biology of the North American Crane Flies (Tipulidse. Dip- 

tera), Part V. Pomona College Jom-. Ent. and Zool., vol. XI. 
pp. 67-74. 

BEYER, ADOLPH. 

214. 1922 — A Brief Resume of Investigations Made in 1913 on Trogoderma 

inclusa. Science Bulletin, vol. XIV (this number). 

BEAMEB, RAYMOND H. 

215. 1916 — An Easy Method of Making Insect Labels. Ent. News, vol. 

XXVIII, p. 418. 

216. 1917— The Oedipodinse of Kansas. Bui. of the Dept.'of Ent., University 

of Kansas, No. 11. pp. 51-126; 74 text figures. 

CLAASSBN, p. \V. 

217. 1914 — Grasshopper Control in the Southern Division of Kansas (with 

Professor Hunter). Jour. Ec. Ent., vol. VII, No. 1, pp. 73-81. 

218. 1917— The Melanopli of Kansas. Bui. of the Dept. of Ent.. University 

of Kansas, No. 11, pp. 5-50; 5 figures. 

CURRAN, C. HOWARD. 

219. 1922— On the Nemestrinid Genus Rhijncocephalus (Nemestrinidse, Dip- 

tera), Can. Ent., vol. LIV. 

220. 1922— Diptera in the Collection of Miss Frances Long (supplement to 

paper by Miss Frances Long on Plant Pollination in Insects) . (In 
press.) 



Historical AccorxT. 13 

DOERINC;. KATHLEEN. 

221. 1922 — Biology and Morpholojiy of Lcpi/rouia qundrangularis (Say). Sci- 

ence Bullf^fiii. \(il. Xn' (this number). 

EMERY, \V. T. 

222. 1914 — Morphologj- and Biologj' of Simulium vittatum and its Distribu- 

tion in Kansas. Science Bulletin VIII; 15 plates. 

H.ACKMAN, LUCTi'. 

223. 1922 — Studies in Cicadella hieroglyphica. Science Bulletin, vol. XIV 

(this number) ; 5 plates. 

HUNGERFOKD, H. B. 

224. 1914 — Anatomy of Simulium vittatum. Science Bulletin, vol. VIII, pp. 

365-382; 3 plates. 

225. 1914 — Notes on Coleoptera from Western Kansas (with F. X. Wilhams). 

Ent. News, vol. XXV, pp. 1-9; 2 plates, 

226. 1915— A Parasite of Cottonwood ■ Borer Beetle. Ent. News, vol. XXVI, 

p. 135. 

227. 1916 — Sciara Maggots Injurious to Potted Plants. Jour. Ec. Ent., vol. 

IX, pp. 538-549; 2 plates, 

228. 1917 — Brief Laboratory Outline for Introductory Entomology. State 

Printer; 18 pages. 

229. 1917 — Egg-laying Habits of a Back Swimmer, Buenoa margaritacea 

Bueno. Ent. News, vol. XXVIII, pp. 174-183; 1 plate. 

230. 1917 — Notes Concerning Food Supply of Some Water Bugs. Science 

N. S., XLV, pp. 336-337. 

231. 1917— Food Habits of Corixids. Jour. N. Y. Ent. Soc, vol. XXV, pp. 1-5; 

1 plate. 

232. 1917 — The Life History of a Back Swimmer, Notonccta undulata Say. 

Ent. News, vol. XXVIII, pp. 267-278, 2 plates. 

233. 1917 — The Life Histoiy of Mesovelia mulsanti White. Psyche, vol. 

XXIV, pp. 73-84; 1 plate. 

234. 1917— The Life History of a Boatman. Jour. N. Y. Ent. Soc, vol. XXV, 

pp. 112-122; 1 plate. 

235. 1918 — Notes on the Ovii)osition of Some Semiaquatic Hemiptera, Jour, 

N, Y. Ent. Soc, vol, XXVI, pp, 12-18; 1 plate. 

236. 1918 — Concerning the Oviposition of the Notonecta. Ent, News, vol. 

XIX, pp. 241-243; 1 plate, 

237. 1919 — Biology and Ecology of Aquatic and Semiaquatic Hemiptera. 

Science Bulletin, vol. XI, pp. 3-328, 33 plates, 

238. 1919 — Male Genitalia as Characters of Specific Value in Certain Crypto- 

cerata. Science Bulletin, vol. XI, pp, 329-332; 2 plates. 

239. 1919 — Biological Notes on Tetradonema plicans Cobb, a Nematode Para- 

site of Sciara coprophila Lint, Jour, of Parasitology, vol. V, pp. 
176-192; 1 plate, 3 text figures. 

240. 1919 — Tables for Determining Types and Breeds of Domesticated Ani- 

mals (third edition). Comstock Publishing Co., Ithaca, N. Y.; 38 
pp., 3 plates. 



14 The University Science Bulletin. 

241. 1920 — Laboratory Outline for Course in Introductorj^ Entomology (re- 

vised and enlarged). World Publishing Co.; 39 pp. 

242. 1922 — Life History of the Toad Bug, Gelastocorus oculatns Fabr. Science 

Bulletin, vol. XIV; 2 plates (this number). 

243. 1922 — A new Subterranean Isopod from Kansas. Science Bulletin, vol. 

XIV; 1 plate (this number). 

244. 1922 — A Review of the Past Quarter Century of Entomology in Kansas 

University. Science Bulletin, vol. XIV (this number). 

245. 1922— The Nepidae of North America. Science Bulletin, vol. XIV; 8 

plates (this number). 

246. 1922 — Oxyhsemoglobin Present in the Back Swimmer, Buenoa margari- 

tacea Bueno. Can. Ent., vol. LIV. 

247. 1922 — Saldoidea slossoni, new var. voileyii. Bui. Brooklyn Ent. Soc, vol. 

XVII, page 64 (Apr.). 

248. 1922 — Notable Additions to Entomological Library at Kansas University. 

Bui. Brooklyn Ent. Soc. (Accepted for publication.) 

249. 1922— Both Hydrometras in Kansas. Bui. Brooklyn Ent. Soc, vol. XVII, 

p. 78 (June). 

250. 1922 — Water Insects from a Portion of the Southern Utah Desert (with 

Dr. R. C. Moore). Science Bulletin, vol. XIV (this number). 

251. 1922— Some Notes on the Egg-laying Habits of the Corixidae. Bui. 

Brooklyn Ent. Soc. (In press.) 

252. 1922 — A Study of the Hydrometra of America North of Mexico, with 

Description of a New Species (Heteroptera, Hydrometridse). Can. 
E'nt., vol. LV. (In press.) 

HUNTER, s. J. 

253. 1914 — University Experiments with Sand Fly and Pellagra. Science Bul- 

letin, vd. VIII. 

254. 1914 — Control Measures for the Native Grasshoppers and Chinch Bugs. 

Bien. Rep. St. Bd. Ag.. vol. XXIV, pp. 680-688; 8 plates. 

255. 1914 — Grasshopper Control in the Southern Division of Kansas (with 

P. W. Claassen). Jour. Ec. Ent., vol. VII, No. 1, pp. 73-83; 5 
plates. 

256. 1915— Report of Official Entomologist. Trans. St. Hort. Soc. of Kan., 

vol. XXXIII, pp. 37-40. 

257. 1915 — Some Economic Results of the Year. Jour. Ec. Ent., vol. VIII, 

No. 2. 

258. 1916— By-products of the Orchard. Country Gentleman, vol. LXXXI. 

259. 1916— Report of the State Entomologist. Rep. Kan. St. Ent. Com. for 

1915-1916, pp. 11-15. 

260. 1917 — Practical Insecticides and Proper Application. Trans. Kan. St. 

Hort. Soc, vol. XXXIV, pp. 182-192. 

261. 1917 — Spring Cankerworm: An Orchard and City Problem. Trans. Kan. 

St. Hort. Soc, vol. XXXIV, pp. 209-212. 

262. 1918 — Municipal Control of Spring Cankerworm. Jour. Ec Ent., vol. XI, 

No. 2, pp. 164-166. 

263. 1918— The Wood Lot. Bien. Rep. Kan. St. Hort. Soc, vol. XXXV, pp. 

144-150. 



HisTOHK AL Account. 15 

264. 1918— Report of tlu> State EntomoloMlist. Rep. Kan. Knt. Coin, for 

1917-191S. i-p. 12-19. 

265. 1918— Report of KntonioioKif't for 1918. Bien. Rep. Kan. St. Hort. Soc, 

vol. XXXV, pp. 171-173. 

266. 1920— Insect Life in Relation to Wheat. Rep. Kan. St. lid. .^ji., vol. 

XXXIX. No. 155, PI). 249-271. 

267. 1921 — Cooperation: Nurseryman and Entoniolotjist . American Nursery- 

man, vol. XXXV, No. 5, pp. 106-107. 

268. 1922— Measures of Prevention, I. Bien. Rep. Kan. St. Hort. Soc, vol. 

XXXVI, i)p. 178-182. 

269. 1922— Measures of Prevention, II. Bien. Rep. Kan. St. Hort. Soc, vol. 

XXXVI, pp. 184-194. 

270. 1922 — The Entomologist and Florist: Their Common Problems. The 

.American Nurseryman, vol. XXXV, No. 5. ])p. 106-107. 

KENNEDY, C. H. 

271. 1917— Dragon Flies of Kansas. Bui. of the Dept. of Ent., University of 

Kansas, No. 11, pp. 127-160; 7 plates. 

L.UVSON, P. B. 

272. 1917 — The Coccidae of Kansas. Bui. of the Dept. of Ent., University of 

Kansas, No. 11, ])p. 161-279; 103 figures. 

273. 1920— The Cicadellida^ of Kansas. Science Bulletin, vol. XII. No. 1, pp. 

1-306; 17 plates. 

274. 1920 — The Cicadidte of Kansas. Science Bulletin, \ol. XII, No. 2, pp. 

306-376; 10 plates. 

275. 1922 — List of the Cicadellidae of Kansas (Homoptera). Trans. Kan. Acad. 

Sci.. vol. XXX, pp. 331-336. 

276. 1922 — List of the Grasses of Douglas County. Trans. Kan. Acad. Sci., 

vol. XXX, pp. 336-339. 

277. 1922— The Genus Aciuopterus (Cicadellidfe). Science Bulletin, vol. XIV 

(this number). 

278. 1922 — The Membracidse of Kansas. Science Bulletin, vol. XIV (this. 

number). 

WELLHOUSE, W.\LTER H. 

279. 1915 — Results of Experiments on the Use of Cyanide of Potassium as an 

Insecticide. Jour. Ec. Ent., vol. IX, No. 1, pp. 169-170. 

280. 1916 — Formulas for Destroying Injurious Insects and Plant Diseases.. 

University of Kansas Circular, No. 4. 

281. 1917 — The Cankervvorm : An Orchard and Shade-tree Pest. Bui. Dept. of 

Ent., University of Kansas, No. 11, pp. 283-315; 3 plates. 

WILEY, GR.ACE OLIVE. 

282. 1922— Life History Notes on Two Species of Saldida>. Science Bulletin, 

vol. XIV (this number). 

283. 1922— Biological Notes on Curictn (Nepida?). Science Bulletin, vol. XIV 

(this number). 

YOUXfi, B. p. 

284. 1918 — Ecological Notes on the Spring Cankerworm (Pulencrlla vernala). 

Can. Ent., vol. L, No. 8, pp. 267-277; figures, 2. 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 3— October, 1922. 

(Whole Series, Vol. XXIV, No. 3.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

The Membr.'Vcid.e of K.ans.as (Homopter.a) P. B. Lawson. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



TABLE OF CONTENTS. 

PAGE 

Introduction 31 

Distribution 32 

Structural characteristics 36 

The male genitalia 38 

Phylogeny of the family 41 

Life history 41 

Economic importance 43 

List of Kansas species 44 

Systematic treatment of Kansas species 45 

Subfamily Centrotin.e 45 

The genus Microcentras 45 

Subfamily Membracin^ ^ 47 

The genus Campylenchia 47 

The genus Enchenopa 48 

Subfamily Smiliin.e 50 

The genus Ceresa 51 

The genus Stictocephala 59 

The genus Acutalis 62 

The genus Micrutalis 63 

The genus Carynota 64 

The genus Thelia 65 

The genus Glossonotus 67 

The genus Heliria 68 

The genus Telamona 69 

The genus Telamonanthc 77 

The genus Archasia 78 

The genus Smilia 79 

The genus Cyrtolobus 80 

The genus Ophiderma 87 

The genus Vanduzea 89 

The genus Entylia 90 

The genus Publilia 91 

Index 107 

(29) 



THE KANSAS UNIYEBSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 3. 



The Membracidae of Kansas. 

By P. B. LAWSON, 

Professor of Eiitoniologj', University of Kansas. 

INTRODUCTION. 

THE writer has been interested in recent years in a systematic 
study of the Homoptera of Kansas. He has previously pub- 
hshed papers on the Coccidse, Cicadelhda^ and Cicadidae of the state, 
listing over 300 species of the members of these families known to 
occur within the borders of Kansas. 

The membracids of the state have been previously studied by 
Miss Hazel Branch, who in 1914 published a paper in the Kansas 
University Science Bulletin, volume 8, on the biology of the 
Membracidse of Kansas. In that paper she listed nineteen species 
from the state. Since that time quite a little collecting has been 
done, until to-day, including some species taken around Kansas 
City, Mo., which species are therefore sure to occur in eastern Kan- 
sas also, there are records of the occurrence of at least fifty-five 
species of tree hoppers in our fauna. Further collecting will of 
course reveal a goodly number of additional species, but the writer 
has thought it advisable to bring our data up to date at this time. 

The writer is greatly indebted to Dr. W. D. Funkhouser, of the 
University of Kentucky, who has made many determinations for 
him and been very generous in giving helpful suggestions and en- 
couragement during the course of this study. Moreover, his paper 
on the biology of the Membracidse of the Cayuga Lake Basin has 
been very freely drawn upon, especially in the use of the technical 
descriptions. 

Through the kindness of Professors Geo. A. Dean and Roger 
Smith, the records of the Kansas State Agricultural College are in- 
cluded in this paper, most of the records from Riley county and sev- 
eral others being from that collection. 

rsi) 



32 The University Science Bulletin. 



DISTRIBUTION. 

The Membracidse, though primarily a tropical and subtropical 
family, are nevertheless found widely scattered and are well repre- 
sented in temperate regions. Authorities are agreed that they are 
best represented in the fauna of Central and South America, but it 
seems certain that when the fauna of Africa and Southern Asia have 
been as carefully studied that these regions will also be found to be 
very rich in these insects. 

Dr. W. D. Funkhouser gives the following distribution of the fam- 
ily according to geographical life zones: 

Palearctic region: 

(Europe, the temperate parts of Asia, and the north of Africa; Iceland and 
the ishinds of the Atlantic; limited by the Himalaj-as.) 

Very poorly represented. Only two or three genera on the entire continent 
of Europe, but two species in Great Britain, two species in Russia, and none 
reported from Iceland. A few in northern Africa, chiefly forms that have 
migrated from the south. 

Ethiopian region: 

(Africa and its islands, except the northern parts; Arabia.) 
Rich in genera and species. Little work has been done on these forms of 
the family, but there is evidence of an abundant membracid fauna. 

Oriental region: 

(India and the East Indies.) 

Extremely rich both in number of forms represented and in number of in- 
dividuals. The center of distribution for the subfamily Centrotinae. 

Australian region: 

(Australia, New Zealand and neighboring islands.) 

Well represented by rather distinct forms. The region has been fairly well 
worked and has yielded a large number of species. 

Nearctic region: 

(America north of Mexico; Greenland). 

Forty or fifty genera, gradually becoming less abundant northward. A 
few species common in Canada as far north as Peny Sound. None reported 
from Greenland. 

Neotropical region: 

(Mexico, West Indies, Central and South America.) 

The most important of all the regions for the Membracidse. Central 
America and the northern part of South America have yielded as many spe- 
cies as all the rest of the world together. 

Altogether over 300 genera have been erected, which contain 
something like 1,500 species. Many of these genera are found in 



Lawson: Membracid.e of Kansas. 33 

more than one of I he abox'c regions, but the number of !>ueh species 
is seemingly rather limited. Thus the American genera Ceresa and 
Stictocephala, while well represented in both the Nearctic and Neo- 
tropical regions, scarcely have a single species that occurs in both 
regions. On the other hand, it is well known that some forms, such 
as Micndalis calva (Say), are found in both the United States and 
the West Indies. 

In North America the members of this family are best represented 
in Mexico. As we advance northward they become fewer and fewer 
till comparatively few species are found in Canada, where they 
reach their northern limit. Van Duzee lists 41 genera in his cata- 
logue of the Hemiptera -of North America north of Mexico. It is 
perhaps safe to say that the number of genera for this region does 
not exceed fifty. The number of species is listed at 185; which com- 
pares very favorably with the 25 cercopids and the 74 cicadas, but 
falls far short of the 357 fulgorids and the 698 cicadellids which oc- 
cur in the same territory. Thus it is seen that in the United States 
this family stands midway in membership among the five families 
of the Homoptera-Aucheinorhynchi. 

So far the membracid fauna of Kansas is known to be repre- 
sented by 20 genera. Further collecting will doubtless reveal the 
presence of several other genera, and of course the number of spe- 
cies will also be increased, for, after all, there has been rather little 
attention paid to this family in the work of past collectors in this 
state. This is shown by the fact that only about 50 out of the 105 
counties of the state have as yet furnished us with specimens. How- 
ever, these counties are so well distributed as to give us a very fair 
idea of the membracid fauna of the entire state. 

The genera Telamona and Cyrtolobus show the most species in 
the state, each having some ten or eleven species. These genera are, 
however, not the ones that are most commonly seen, for both 
Stictocephala and Ceresa are far more abundant as to individuals, 
Ceresa bubalus (Fabr.) being by far the commonest species in the 
state. Close to the two latter genera comes Micrutalis, the species 
calva having been taken in about fifteen counties. Campy lenchia 
latipes is also very common throughout the alfalfa fields of the 
state. Vanduzea triguttata (Burm.) occurs in enormous numbers 
when taken, as does Entylia concisa (Walk.), but hitherto these 
species have not been taken in many places within our borders. 



34 The University Science Bulletin. 



EXPLANATION OF PLATE I. 

1. Pronotum and head of Ceresa bubalus. sp, suprahuraerals ; m, meto- 
pidium; ha, humeral angles. 

2. Cephalic aspect of head of Ceresa bubahis. es, epicranial suture; o, 
ocellus; v, vertex; e, compound eye; g, gena; lo, lora; cl, clypeus; I, labrum; 
ep, epipharynx; la, labium. 

3. Caudal aspect of head of Cerem bubalus. oc, occiput; of, occipital 
foramen; pg, postgena; c, compound eye; la, labium. 

4. Lateral aspect of head of Ceresa bubalus. oc, occiput; v, vertex; e, 
compound eye; g, gena; cl, clypeus; pg, postgena; I, labrum; ep, epipharynx; 
mx, maxillary stylets; md, mandibular stylets; la, labium. 

5. Pronotum of Ceresa bubahis. sp, suprahumerals; ha, humeral angles; 
m, metopidium; dc, dorsal carina; pp, posterior process. 

6. Metathoracic leg of Ceresa bubalujs. c, coxa; tr, trochanter; /, femur; 
t, tibia; ta, tarsus. 

7. Prothoracic leg of Ceresa bubalus. 

8. Prothoracic leg of Campylenchia latipes. 

9. Tegmen of Ceresa bubalus. 
10. Hind wing of Ceresa bubalus. 

IL Ventral aspect of abdomen of male Ceresa bubalus. 2-9, sternites; 2-8, 
pleurites; py, p.ygofer or ninth tergite; Iv, lateral valve. 

12. Ventral aspect of abdomen of female Ceresa bubalixs. 2-7, sternites; 
2-S, pleurites; pif, pygofer or ninth tergite; ovi, ovipositor. 

13. Lateral aspect of abdomen of male Ceresa bubalus. 1-11, tergites; 2-9, 
sternites; Iv, lateral. valve. 

14. Lateral aspect of abdomen of female Ceresa bubalus. 1-9, tergites; 2-7, 
sternites. 



Lawson: ISIembracid.e of Kansas. 



35 



PLATE I. 




36 The University Science Bulletin. 



STRUCTURAL CHARACTERISTICS. 

The outstanding characteristic of the membracids is their re- 
markably developed pronotum. This sclerite is usually enlarged 
so as to cover most of the thorax and much of the abdomen, and 
while in our species it is often extended into rather regularly formed 
processes, yet in many tropical species these processes assume the 
most irregular and grotesque shapes. In the subfamily Centrotinse, 
however, the pronotum is not thus enlarged, but is so small that the 
greater part of the scutellum is left exposed. 

As in all insects, the body is divided into three regions. The head 
is quite characteristically homopterous, fitting against the coxae of 
the prothoracic legs, between which lies the beak. At its lateral 
extremities are the well-developed compound eyes, while the simple 
eyes or ocelli are found on the i ertex, the paired sclerite forming 
the greater part of the cephalic aspect of, the head. The ocelli are 
always paired and in a line with each other and the compound eyes, 
though their distance from each other and from the compound eyes 
varies. 

The setaceous antennce are found under the margin of the vertex 
on either side of the clypeus. The first three segments are large, but 
the rest of the organ is composed of a large number of small 
segments which decrease in size toward the apex. 

The vertex is a paired sclerite lying on either side of the base of 
the Y-shaped epicranial suture. The presence of this suture would 
indicate a mlore primitive position for this family than for the 
Cicadellidae, where no signs of it are present. 

Between the arms of the epicranial suture and forming the apex 
of the head is the clypeus. This sclerite is often characteristic in 
the length of its apex and in its relation to the lateral margins of 
the vertex. At its apex it is turned backward, forming a rounded 
lobe, caudad of which are the genoi. The latter sclerites are seen 
to form the lateral portions of the head, extending between the eyes 
and the clypeus, and bearing the antennae. The postgence occupy 
most of the caudal aspect of the head, forming, with the dorsal 
occipital sclerites or occiput, most of the boundary of the occipital 
foramen. The lorce are found on either side of the clypeus and 
labrum as two more or less distinct sclerites with rounded lateral 
margins. 

The beak or labium consists of three segments, as in other 
Homoptera, and contains in a groo^'e the mandibular and maxillary 



Lawson: Membracid^ of Kansas. 37 

stylets. The labrum is large aiul cauchul of the elypeus, bearing the 
more or less membranous epipharynx at its tip. All the mouth 
parts are typical of the ordinary sucking phytophagous insect. 

The thorax, as in all insects, consists of three segments. For our 
purposes only a brief discussion of the dorsal sclerite of the pro- 
thorax will be necessary. This sclerite, the pronotum, as already 
mentioned, is the chief characteristic of the family. It is one of 
the finest examples of the biological phenomenon of orthogenesis 
that is to be found. In our North American membracids it is com- 
paratively simple as compared with its appearance in many Neo- 
tropical genera, such as Sphongophorus and Cyphonia. But even in 
our fauna it assumes such diverse forms as to give us the best 
generic, and often also specific characters for the classification of the 
members of this family. The following parts have been named and 
are used in the descriptions which follow: 

1. Metopidium. The cephalic area ,of the pronotum, extending 
from the base of the head to the front of the dorsum. 

2. Humeral angles. The angles of the pronotum just above the 
bases of the wings. 

3. Suprahumerals. The lateral projections above the humeral 
angles. 

4. Dorsal carina. The ridge usually extending the length of the 
pronotum. 

5. Posterior process. The caudally directed portion of the 
pronotum. 

The three segments of the thorax each bear a pair of legs, which 
are composed of the usual segments, the tarsi being three-segmented. 
The coxce are usually stout, the trochanters rather small and bent, 
the femora large and stout, and the tibice usually long, slender and 
hairy. In the subfamily Membracinae and in some Centrotinae, the 
tibiae are wide and flat or foliaceous, especially so in the prothoracic 
and mesothoracic legs. 

The last two thoracic segments each bear a pair of wings, which 
are of value in sj^steraatic work, not only in their venation, but also 
in their relation to the pronotum. 

The abdomen consists of eleven segments, the last two forming 
the anal tube. Most of these segments show a distinct tergite, 
pleurites and sternite. Of these sclerites the tergite is by far the 
largest, forming both the dorsal and lateral portions of the seg- 
ment; and in the case of the female ninth segment, nearly inclosing 
the segment, coming clear down to the sides of the ovipositor, in 



38 The University Science Bulletin. 

this sex there showing externally but six sternites, those of the 
second to seventh segments, inclusive. In the male the ninth tergite 
is not as large, for in this sex the sternites of segments two to nine, 
inclusive, all show clearly. This tergite bears, however, on either 
side, a pair of broad lobes or plates which Funkhouser calls the 
lateral valves. These, he thinks, may be the pleura of the ninth 
segment and of use in copulation. They may project caudad or 
turn mesad — a character of generic value. They frequently bear 
variously shaped processes or teeth, which, as far as the writer has 
observed, are constant in form and position for the various species, 
and are therefore good specific characters. 

The sternite of the seventh segment of the female differs in differ- 
ent species, and is therefore of systematic value, as is also the 
ninth sternite or sternal plate of the male. The latter is constant 
in shape and extent of apical splitting within the species, and this 
makes it of value as a taxonomic character. 

THE MALE GENITALIA. 

As in the Cicadellidae, the abdomen of the male ends in what 
Sharp calls the "terminal chamber" in the case of the Pentatomidse. 
This chamber is bounded above by the anal tube and ninth tergite, 
laterally by the lateral valves, and ventrally by the ninth sternum. 
Within this chamber are found the genital organs of the male, their 
position and structure being exactly homologous with the male 
genitalia of the leaf hoppers. 

The styles or claspers are always paired, both members of the pair 
being alike, and fastened to the sternal plate by apparently passing, 
near their middle, through the membrane forming the dorsal surface 
of the plate. The cephalic portion of the clasper projects into the 
abdominal cavity, frequently reaching the sixth segment, though 
usually extending only as far as the eighth or ninth. It does not 
seem to vary much in the different species. The apical portion, 
however, is characteristic in its shape and apical structure, some- 
times being nearly straight, at other times more or less strongly 
curved, and ending either in a plain acute point or in variously 
shaped and toothed extremities. So far the writer has found no two 
species in w'hich the styles are alike in both size and structure. 

Canon Fowler, in the "Biologia Centrali-Americana," states that 
in the genus Ceresa the styles are long and pointed, while in the 
genus Stictocephala they are short and obtuse at the apex. This 
is undoubtedly frequently true, but the writer has found that it does 



Lawson: Membracid.e of Kansas. 39 

not always hold in tlu' members of these genera as the latter are now 
constituted. Thus, while in Stictocephala festina the above-men- 
tioned condition holds good, it is not true of S. lutea, for the latter 
possesses styles just as long and pointed as in several members of 
Ceresa, notably C. bubalus. 

As in the Cicadellida>, the stvles are united bv a small sclerite, 
whicii \-aries in shape in the various genera and species, but is 
always of the same shape within the species. This sclerite the 
writer has called the "style-oedagus connective," or more simply, 
the "connective.'' The former name shows its usual function in the 
leaf hoppers, for usually it unites not only the two styles, but at its 
distal end is united to the base of the cedagus. This condition pre- 
vails in all the tree hoppers thus far examined. Its relation to the 
styles, however, is apparently different in the two families, for in 
the leaf hoppers each style sends out a distinct chitinous process to 
which it is fastened, while in the tree hoppers this chitinous process 
is not usually present, but rather each style, at the point of attach- 
ment to the connective, gives off a dorsal membranous fold which 
unites with the connective, the two folds appearing heart-shaped 
basally when viewed from above. 

In all the species so far examined the connective appears as a 
rather thin, usually fiat sclerite, frequently showing a distinct tend- 
ency to fold longitudinally along a median keel, thus drawing the 
two styles closer together. It is usually quite small, but in some 
forms, notably Ceresa borealis, it is well developed. 

The writer has not tried to determine the morphological status of 
the connective. Kornhauser calls it the ninth sternite. Funkhouser 
does not mention it, but feels that the last sternal plate or the valves 
are the ninth sternite. That it is a sternite cannot be doubted, and 
the writer hopes to discuss its exact position in a later paper on the 
genitalia of the Homoptera-Auchenorhynchi. 

The cedagus, or perns sheath, seen laterally, is usually a V-shaped 
organ, the mouth of the V looking caudad or dorsad. The anterior 
or dorsal arm extends up to the base of the anal tube, the latter 
always being fastened to it, usually at a slight excision near its 
apex. As far as our present studies go, the shape of this arm seems 
to vary more or less even wathin the species, sometimes being rather 
narrow and straight, then changing till it is frequently quite broad 
and variously bent. The lower or posterior arm is the penis sheath 
proper. The penis enters it just above the usual basal angle by 
which the cedagus is united to the connective. This basal angle may 



40 The University Science Bulletin. 

often be prolonged into a distinct cephalic process. This ventral arm 
varies greatly in different genera and species, but, as far as our 
observations go, is always constant in shape within the species. It 
usually extends more or less dorso-caudad, though sometimes its 
apex is bent distinctly dorsad and may even extend dorso-cephalad. 
The course of the penis through this arm is usually easy to trace in 
caustic potash specimens. 

In the genera Ceresa and Stictocephala the functional orifice is 
usually along the ventral side of this arm and at quite a distance 
from the apex. This opening in these genera is usually quite long 
and is guarded by a distinct membranous external sheath. In S: 
jestina, however, the penis opens apically, while in Campylenchia, 
Telamona and others it opens at or near the apex. In every case, 
however, the opening seems to be guarded by the external membrane 
and is constant in position within the species. The shape of the 
apex also is constant for the species ; in fact, the whole organ seems 
to afford an excellent specific character for use in systematic work. 
Canon Fowler's characterization of the cedagus of Stictocephala 
does not hold good for all the species of the genus. 

The cephalic portion of the styles and the dorsal arm of the 
cedagus are practically internal parts. It seems as though these 
parts have distinct layers added to them at various intervals, thus 
adding to the length of the former and to the width of the latter 
This may account for the variable condition seen in these organs, 
even within the species. The terminal portions, however, seem to 
be constant. 

The sternal plate, both as to shape and the extent of its division 
into two parts, is an excellent and readily available character for 
taxonomic work. It may be the ninth sternite, but the question as to 
its possible homology with abdominal appendages at once arises. 
At any rate, it seems to be exactly homologous with the plates of the 
Cicadellidse. 

Further work with these genital organs will undoubtedly lead to 
their greater use in systematic work. The writer has found that 
specimens identified as identical can frequently be separated and 
properly classified by a study of these organs. It is to be hoped that 
the various genera will be studied from this standpoint in the days 
to come. If they are the writer feels that several generic changes 
will be made. Thus it would seem that the genus Stictocephala 
should be divided, for the genitalia of S. festina and S. lutea cannot 
possibly belong to members of the same genus. On the other hand, 



Lawson: Membraoid.t: of Kansas. 



41 



the genitalia of the genera Tclamona and Tclonaca are so near alike 
that it seems certain that the latter should never have been separated 
from the formei-. 

PHYLOGENY OF THE FAMILY. 

The membracids belong to that group of the Homoptera known 
as the Auchenorhynchi, which differs from the rest of the members 
of the order, the Sternorhynchi, in that in the latter the mouth parts 
seeminglj'- arise from between the prothoracic legs, instead of from 
the head, as in the former. 

In this group are five families, arising from three different stems, 
of which it seems clear that the Cicadidae arise from the lowest stem 
and the Fulgoridse from the highest. In between these two families 
are the closely related membracids, cicadellids and cercopids. As 
pointed out in his paper on the Cicadellidse of Kansas, the writer 
accepts the conclusions of Funkhouser and others in making the 
membracids the lowest of these three families and the cercopids the 
highest. This relationship would therefore be expressed as follows: 



FULGORIDAE 




CERCOPIDAE 



CICADELLIDAE 



MEMBRACIDAE 



CICADIOAE 



In the above figure it will be noticed that the Cercopidae are rep- 
resented as branching off earlier from the median stem than the other 
two families. This seems evident when their life history is consid- 
ered, for they have been isolated long enough to enable them to per- 
fect a method of protection against parasitism. Certainly the devel- 
opment of the production of the spittle mass which incloses the 
nymph could not have occurred quickly, but must have taken a 
relatively long period of time to attain its present perfection. 

LIFE HISTORY. 

Most of the members of this family occur on small trees or shrubs, 
particularly those growing near the edges of woods or out in the 
open. In the main they seem to avoid the shade and prefer those 



42 The University Science Bulletin. 

situations where they receive the most sunshine. Most of our 
species occur on trees, but there are several that are found on 
shrubs^ and some occur on weeds, grasses and clovers. 

The majority of our species of tree hoppers are found to over- 
winter in the egg stage. This stage, therefore, occupies by far 
the longest period in the life history of the insect. The eggs are 
deposited for the most part in the twigs. Here they may be placed 
in simple incisions in a row, or the well-known method of oviposi- 
tion of Ceresa biibalus may be used where two curved iuQisions, 
facing each other, are made. Some species also oviposit in the 
buds, while our common Entylia concisa lays its eggs in the veins 
of the leaf of Ambrosia trifida. Funkhouser gives the axils of the 
leaves as the place where Telamona ampelopsidis oviposits, while 
Thelia bimaculata lays most of its eggs in the roots or on the stem 
below the surface of the soil. 

While most of our treehoppers hibernate in the egg stage, Funk- 
houser states that Entylia bactriana and Publilia concava over- 
winter as adults. The writer has taken Entylia concisa late in the 
fall in the rubbish and grass around its host plant. In fact, on 
September 8 he has found the eggs of this species just hatching, 
while on September 11 he has taken all the nymphal instars and 
newly matured adults on a single leaf. 

The number of generations in a season is usually one, by far the 
larger number of our species having a single annual brood. Some 
species, however, have two broods and some probably three, while 
Vanduzea arquata is said by Funkhouser to have as many as four 
generations. The same is seemingly true of our Vanduzea trigut- 
tata. 

The usual life history of a tree hopper has been summarized by 
Doctor Funkhouser as follows: 

Eggs: Laid in fall, hatch in early spring. 

Nymphs: Emerge about the middle of May and require about six weeks 
to reach maturity. 

Adults: Are common about July 1 and persist throughout summer and fall. 

Mating: Takes place the first week after emergence. 

Oviposition : Occui"s within a week after mating. 

Broods: Usually one but sometimes more, dependent on weather condi- 
tions. 



Lawson: Membkacid.f. of Kansas. 43 

For a single individual the life cycle would be somewhat as follows: 

Egg stage: From SenleiiihiT to middle of May 8V^ months 

First instar 1 week 

Second instar 1 week 

Third instar 1 week 

Fourth instar 1 week 

Fifth instar 2 weeks 

Nymph : 

Total, fniiii middle of May to July 1% months 

Adult : From July to October (inclusive) 4 months 

Entire life 14 months 

The above dates of course apply to New York and would differ 
somewhat from what occurs in Kansas. 

One of the easiest ways to find certain species of tree hoppers is 
to notice the more readily seen ants which attend so many of our 
common forms. In this way Vanduzea triguttata and Entylia con- 
cisa are very easily located. It is well known that the ants obtain 
honey dew from these insects, although there are many species of 
tree hoppers which do not seem to furnish this substance in suffi- 
cient ciuantity to attract the attention of these visitors. 

It is well known that these insects are frequently parasitized. 
Hymeqopterous parasites of the family Dryinidae have been known 
to occur in several genera of tree hoppers. Doctor Kornhauser 
worked out in great detail the life history of such a parasite, 
Aphelopiis thelice, which parasitizes Thelia himaculata. In addi- 
tion, tree hoppers are often the prey of predaceous insects and 
spiders. Of such predaceous enemies there are a number of kinds, 
but it is quite doubtful if these enemies are a very vital factor in 
the natural control of the members of this family. 

ECONOMIC IMPORTANCE. 

The chief damage done by tree hoppers is by their oviposition. 
The records of injury to young orchards, especially to apple, are 
many. The type of oviposition that may really be serious is that 
represented by the egg punctures of Ceresa bubalus, and this species 
is seemingly the most often accused of all the species concerned. 
The writer has seen young orchards in which the outer twigs were 
all roughened up by the oviposition scars, and such cases are numer- 
ous in the literature. The exact damage done, however, is prob- 
lematical, many inclining to the opinion that as a rule the damage 
done is not serious. Others hold that in after years such scars will 
weaken the branches, causing them to be more easily broken off by 
the wind, and also furnishing good points of attack for borers. The 



44 



The University Science Bulletin. 



wounds made also doubtless furnish good points of entrance for cer- 
tain fungous diseases. 

Cases are on record also where the buds of certain trees were 
seriously injured by the deposition of eggs within them. Damage 
from such oviposition always results when eggs are deposited deeply 
in small buds. In this respect the species concerned most fre- 
quently are Ceresa taurina, Stictocephala inermis and Enchenopa 
binotata. The species most injurious to stems belong to the genera 
Ceresa and Stictocephala. 

Inasmuch as the nymphs of the most injurious species prefer to 
feed on the more succulent weeds and grasses in the orchard, the 
clean culture of badly infested orchards and the getting rid of 
near-by weed patches in the early summer, will usually be all the 
control measures necessary. Frequently in case of very severe in- 
festations the careful pruning of the worst -infested twigs after egg 
deposition is a desirable practice. 



LIST OF THE SPECIES. 



Microceniru>i caryce (Fitch). 
Campylenchia latipes (Say). 
Enchenopa binotata (Say). 
Ceresa diceros (Say). 
Ceresa albescens Van D. 
Ceresa bubalus (Fabr.) 
Ceresa taurina Fitch. 
Ceresa palmeri Van D. 
Ceresa borealis Fairm. 
Ceresa brevicornis Fitch. 
Stictocephala inermis (Fabr.) 
Stictocephala festina (Say). 
Stictocephala lutea (Walk.). 
Acutalis tartarea (Say). 
Micrutalis calva (Say). 
Carynota mera (Say). 
Thelia bimacnlata (Fabr.). 
Thelia uhleri Stal. 
Glossonotus acuminatus (Fabr.) 
Heliria cristata (Fairm.). 
Heliria s^alaris (Fairm.). 
Telamona pyramidata (Uhl.). 
Telamona viridia Ball. 
Telamona obsoleta Ball. 
Telamona decorata Ball. 
Telamona querci Fitch. 
Telamona ampelopsidis (Harr.). 



Telamona tristis Fitch. 
Telamona unicolor Fitch. 
Telamona extrema Ball. 
Telamona collina (Walk.). 
Telamonanthe rileyi (Godg.). 
Telamonanthe modesta (Godg.). 
Archasia galeata (Fabr.). 
Archasia belfragei Stal. 
Smilia camelus (Fabr.). 
Cyrtolobus celsus Van D. 
Cyrtolobus juliginosus (Emns.). 
Cyrtolobus arcuatus (Emns.). 
Cyrtolobus jenestratus (Fitch). 
Cyrtolobiis tuberosus (Fairm.). 
Cyrtolobus I'au (Say). 
Cyrtolobus griseus Van D. 
Cyrtolobus cinereus (Emns.). 
Cyrtolobus querci (Fitch). 
Cyrtolobus muticus (Fabr.). 
Ophid^rma salamandra Fairm. 
Ophiderma flaviguttida Godg. 
Ophiderma flava Godg. 
Vanduzea triguttata (Burm.). 
Entylia concisa Walk. 
Pnblilia concava (Say). 
Publilia modesta Uhl. 
Publilia reticulata Uhl. 



Lawsox: Membracid.T': of Kansas. 45 



SYSTEMATIC TREATMENT OF KANSAS SPECIES. 

The three subfamilies represented in Kansas may be separated 
by the following key : 

KEY TO SUBFAMILIES. 

A. Scutelhun distinct, not concealed by the pronotum. Centrotinse. 
AA. Sciitelhini wanting or concealed by the pronotum. 

B. Tibiae, especially anterior ones, foliaceoiis. Membracinae. 

BB. Tibiae not foliaceoiis. Smiliinae. 

Subfamily CENTROTIN^ (Spinola). 

This subfamily, represented in the United States by about half a 
dozen genera and a few more species, is readily separated from the 
other subfamilies by the presence of a usually visible scutellum. 
The genus Microccntrus occurs in Kansas. 

Genus Microcentrus (Stal). 

The members of this genus that occur in the United States have 
a pronotum that leaves a large part of the scutellum well exposed 
and ends in a slender terminal process which extends beyond the 
divided apex of the scutellum. 

Both members of this genus known to occur in the United States 
have been taken in Kansas. They may be separated by the follow- 
ing key : 

KEY TO SPECIES. 

A. Prothorax bearing a pair of laterally flattened horns. perdita. 

AA. Prothorax without laterally flattened horns. caryce. 

Microcentrus caryce (Fitch). 

Uroxiphus caryce Fitch, Hornop. N. Y. St. Cab., p. 52, 1851. 
Centrotus caryce Walker, List Homop., iv, p. 1147, 1851. 
Microcentrus caryce Stal, Of. Vet. Akad. Forh., xvi, p. 295, 1869. 
Piiaulocentrus caryce Fowler, Biol. Centr. Am., Homop., ii, p. 159, 1896. 

Funkhouser gives the following technical description: 

Gray-brown mottled with black; entire body broad and flat; pronotum 
roundly swollen above line of abdomen and wings; wings broadJy tectiform. 

Head perpendicular, twice as broad as long, roughly sculptured, closely jnmc- 
tate, densely pubescent, deep brown at base; eyes prominent, extending be- 
yond lateral margins of pronotum, dark brown margined with lighter; ocelli 
small, pearly, farther from the eyes than from each other, with deep depression 
between them; clypeus prominent, broad, lighter in color than vertex above, 
extending far below lateral margin of head. 

Prothorax subspherical, with high median carina, coarsely punctate, pubes- 
cent ; light brown marked with black on median ridge and above head ; pos- 
terior margin tnmcate except for narrow process, which projects to angles of 



46 The University Science Bulletin. 

tegmina and short sharp tooth on each latero-posterior angle. Scutellum 
broadly exposed, wide at base, truncate at tip, which does not reach apex of 
posterior process. 

Tegmina translucent, pubescent, inner margins straight and meeting at 
median dorsal line; veins prominent and nodulate; apices of tegmina extend- 
ing beyond tip of abdomen. Legs and undersurface of body light brown 
mottled with white. Undersurface of abdomen often tomentose. 

Length, 9-10 mm.; with, 3 mm. 

Distribution. Van Duzee records this species from Ontario, New 
York, New Jersey, Pennsylvania, North Carolina and Ohio. In the 
Crevecoeur collection is a specimen taken in Pottawatomie county, 
Kansas, thus establishing its occurrence in the state. 

Hosts. As indicated by the specific name, this species occurs on 
hickory. Funkhouser reports it on oak also. Matausch gives swamp 
oak and walnut, and Goding mentions the latter. 

Microcentrus perdita (Amyot & Serville). 

Ledra perdita Amyot & Serville, Heinip., p. 577, pi. 11, fig. 5, 1843. 
Centruchus leibecki Goding, 111. St. Lab. Nat. Hist., iii, p. 471, 1894. 
Microcentrus perdita Baker, Can. Ent., xxix, p. 38, 1897. 
Centruclioides perdita Van Duzee, Bui. Buf. Soc. Nat. Sci., ix, p. 117, 1908. 

Goding describes this species as follows: 

Yellow-ferruginous, silky white between lateral horns; behind horns a spot 
on costal margin, near base of tegmina. Head black, broad, eyes verj' prom- 
inent; base convex, griseous, lower part of face strongly declivous; four rough- 
ened caringe pass along face from base do\vnward, the ocelli being in the two 
internal ones, the outer ones being contiguous to the ej'es; part of face below 
eyes triangular, apex yellow. Ocelli equidistant from each other and the eyes. 
Prothorax convex, lateral angles slightly produced, a prominent median carina 
extending from base to apex nearly black. Above the lateral angles, on each 
side, is a long horn or protuberance, flattened laterally, slightly curving upward, 
outward and forward, the apex truncated; width of base and apex equal. Apex 
of scutellum bidentate, the teeth ivory-white; posterior margin of prothorax 
with a very slender tooth or style, extending backward on each side of pos- 
terior process of prothorax, a little distant from it. The entire surface of the 
prothorax densely and regularly punctured. Apex of tegmina far surpassing 
tip of abdomen; a black spot on the internal margin a short distance from the 
apex; another black spot on the costa, about one-third the distance from the 
base. Tarsi black; legs mottled with ferruginous and grayish-yellow; tibiae 
triquetrous. Tegmina lightly ferruginous and opaque. 

Length to apex of tegmina, 8.5: mm.; width at lateral angles, 3.2 mm. 

Distribution. Reported from Massachusetts, New Jersey, Penn- 
sylvania, Florida, Colorado and New Mexico. Specimens have also 
been taken in Riley county, Kansas. 

Hosts. Van Duzee reports this specief> from scrub oak. 



i 



Lawson: Membracid.e of Kansas. 47 

Si-BFAMTLY I\I KMBRACIX.E (Stal). 

The members of this subfamily differ from all other mcmbracids 
of our fauna by having the prothoracic and mesothoracic tibiae 
foliaceous. As in the following subfamily, the pronotum conceals 
the scutellum. 

Two genera occur in Kansas ^^■ilich may be separated by the fol- 
lowing key: 

KEY TO GEiN'ERA. 

A. Lateral ridges of anterior horn n(\irer dorsal carina; ventral carina of horn 
not distinctly foliaceous. Campylenchia. 

AA. Lateral ridges of anterior horn midway between dorsal and ventral 
carinse; latter distinctly foliaceous. Enchenopa. 

Genus Campylenchia (Stall. 

This genus, the chief characteristics of which appear in tlie above 
key, is represented by a single though widely spread species in the 
United States. 

Campylenchia latipes (Say). 

(PI. VI, figs, 9, 10.) 

Membracis latipes Say, Long's 2nd Exped., ii, p. 302, 1824; Conipl. Writ., i, p. 202. 

Enchophylluvi latipes Fitch, Homop. N. Y. St. Cab., p. 47, 1851. 

Enchenopa latipes Walker, List Homop., ii, p. 482, 1851. 

Enchenopa antonina Walker, List Homop., ii, p. 488, 1851. 

Enchenopa venosa Walker, List Homop., ii, p. 488, 1851. 

Enchenopa frigida Walker, List Homop., ii, p. 490, 1851. 

Enchenopa himacula Walker, List Homop., ii, p. 491, 1851. 

Campylenchia curvata Stal, Heniip. Fabr., ii, p. 43, 1869. 

Enclienopa curvata Uhler, Bui. U. S. Geol. Geog. Surv., i, p. 343, 1876; iii, p. 457, 1877. 

Aconophora curvata Butler, Cist. Ent., ii, p. 349, 1877. 

Echenopa rectidorsian Buckton, Monog. Membr., p. 49, 1903. 

Campylenchia latipes Van Duzee, Can. Ent., xliv, p. 320, 1912. 

Funkhouser gives the following technical description: 

Uniform cinnamon brown, densely punctate, sparingly pubescent; single 
porrect pronotal horn projecting forward over head; head and first two pairs 
of legs broadly foliaceous, hind legs spined; tegmina opaque, punctate at basal 
and costal margins. 

Head quadrate, somewhat declined, shining brown somewhat mottled with 
darker, lightly punctate, densely pubsecent ; eyes prominent; ocelli small, 
pearly, equidistant from each other and from the ej'es and situated on a line 
passing through centers of eyes; clypeus very broad, shining, scarcely punctate, 
broadly truncate at apex, tip strongly pubescent. 

Prothorax produced anteriorly into a long, flattened horn, ridged in center 
and foliaceous above and below, varying greatly in length and degree of curve; 
posterior process strong, tectiform, reaching internal angles of tegmina; 
median dorsal carina strong and percuiTent ; entire pronotum concolorous, 
lightly punctate, sparingly pubescent with golden hairs; median lateral ridge 
reaching lateral margin. 

Tegmina yellow-opaque; basal and costal areas punctate and pubescent; 



48 The University Science Bulletin. 

veins distinct, broad, and slightly pubescent ; fi\e apical and two discoidal 
cells; hind wings iridescent. Two anterior pairs of legs broadly spatulate and 
lightly pubescent at margins; posterior tibiae armed with black-tipped spines; 
tarsi much produced and lighter in color. Undersurface of body chocolate 
brown. 

Length: from head to apices of elytra, 5 mm.; from tip of promotal horn 
to apices of elytra, 8 mm. Width between humeral angles, 2 mm. 

Internal male genitalia. Styles with long flattened anterior proc- 
ess, widest apically, with a distinct prominence at attachment to 
connective, caudad of which they are peculiarly wrinkled and show 
a small but distinct mesal lobe, then widest just caudad of wrinkled 
portion, after which they curve laterad, ending in a doubly pointed 
apex, of which the terminal tooth is larger and more slender than 
the anterior one, the apical third of the styles bearing a few short 
hairs; connective short and wide, basal half with a heavily chitinized 
and flattened V-shaped portion; oedagus, viewed laterally, stout and 
broadly V-shaped, widest at the distinctly notched base, the upper 
basal process also slightly notched, narrowest at the bend and then 
widening till near the apex, the caudo-dorsal portion of which is only 
slightly chitinized, the functional orifice being at about the middle 
of this slightly chitinized area, the upper portion of the apex bearing 
a few distinct file-like teeth. 

Distribution. Van Duzee gives the following distribution for this 
species: Ontario, Maine, Vermont, New York, New Jersey, Penn- 
sylvania, North Carolina, Illinois, Iowa, Kansas, Missouri, Colo- 
rado, Wyoming, Montana, Texas, New Mexico, Arizona, and Cali- 
fornia. Osborn reports it from Florida. It is thus seen to extend 
throughout the United States. 

In Kansas specimens have been taken in the following counties: 
Douglas, Ottawa, Pratt, Hodgeman, Clark, Pottawatomie, Labette, 
Bourbon, Wabaunsee, Russell. Riley, Ellis, Ellsworth and Wyan- 
dotte. 

Hosts. Funkhouser gives the following hosts for this species: 
Aster, daisy, joe-pye weed, alfalfa, prickly lettuce, wild carrot. 
Miss Branch mentions golden rod and sensitive rose. Gillette and 
Baker report it from Glycyrrhiza lepidota, Psoralea tenuijiora and 
Apocynimi androsacniifolium. The writer has found it very abun- 
dantly on alfalfa. 

Genus Enchenopa Amyot & Serville. 
But two members of this genus are known to occur in the United 
States. The widely distributed E. binotata occurs in Kansas. 



i 



Lawson: Membracid.e of Kansas. 49 

Knchcnopa binotnfa (Say). 
(I'l. \i, ti«s. n, 12.) 

Moiibracis biiiotata Say, Long's 2nd E.\ped., ii, p. 3(ll, 1824; Conipl. Wril., 1, p. 201. 

Enchophyllum binotatum Fitch, Homop. N. Y. St. Cab., p. 47, 18.51. 

Enchenopa biiiotata Walker, I.,ist Homop., ii, p. 481, 1851. 

Encheiiopa brevis Walker, List Homop., ii, p. 492, 1851. 

Thelia biiiotata Emmons. Nat. Hist. N. Y. Ins., p. 15f>, 1854. 

Enchenopa bifusifera Walker, List Homop., f<uppl., p. 125, 1858. 

Enchenopa bivittata Rathvon in Mombert's Hist. Lane. Co., Pa., p. 551, 1869. 

Enchenopa porrecta Buckton, Monog. Membr., p. 51, pi. (i, fig. 5, 1902. 

Funkhouser gives the following technical description: 

Much resembling the preceding species in size and in general appearance, 
but differing in shape of the head, in shape of sculpturing of the pronotal 
horn, and in bearing two yellow spots on the dorsal line of the pronotum. 

Head longer than broad, uniform brown, finely but densely punctate, spar- 
ingly pubescent; eyes prominent, very deep brown; ocelli yellowish, farther 
from each other than from the eyes; clypeus longer than broad, rounded at 
tip, not punctate. 

Prothorax finel\- i)unctate, sparsely pubescent; two distinct ridges on each 
.side, the upper extending to the lateral margin; pronotal horn strongly curved, 
broadly foliaceous above, triquerate at tip; median dorsal carina high and per- 
ciu-rent; two dorsal spots of lemon yellow, the anterior about twice as long 
as the posterior; posterior process gradually acuminate, extending slightly 
beyond internal angles of tegmina. 

Tegmina concolorous brown, opaque, costal margins slightly jjunctate, and 
feebly pubescent at base; veins distinct; five apical and one discoidal cell. 
First two pairs of legs broadly fohaceous; hind tibiae spined; tarsi thin. 

Length 5 mm.; width 2 mm. 

Internal male genitalia. Styles stout, anterior portion longer and 
broader than posterior, with a rounded prominence to connective, 
wider just caudad of this, apices strongly hooked and truncate, the 
apical point being slightly longer than the anterior one, with a few 
hairs on the terminal third; connective heart-shaped, broad, a basal 
V-shaped band heavily chitinized; cedagus, viewed laterally, 
strongly curved, base very wide and extending dorsad in a large 
rounded prominence which is separated by a broad notch from the 
pointed attachment to the connective, the apex pointed and with 
functional orifice below the tip, the dorsal surface of which bears 
a number of filelike teeth. 

Distribution. Van Duzee reports this species from Ontario, Mas- 
sachusetts, New Hampshire, New York, New Jersey, Pennsylvania, 
Maryland, District of Columbia, North Carolina, Florida, Illinois, 
Iowa, Michigan, Kansas, Missouri and Texas. In Kansas speci- 
mens are reported from Douglas, Riley, Sedgwick, Bourbon and 
Pottawatomie counties. 



50 The University Science Bulletin. 

Fowler records it from Mexico, Guatemala and Panama. It is 
thus seen to occur throughout tlie eastern United States and down 
through Central America. 

Hosts. Funkhouser gives the following hosts: Locust, wild grape, 
bittersweet, hickory, sycamore, butternut, dogwood, daisy, joe-pye 
weed. Miss Branch adds golden rod and pin oak. Matausch gives 
Viburnum. Goding records it from butternut, birch, apple, walnut, 
grape, hop tree, locust, redbud, cherry. Viburnum, Ceanothus and 
white birch. The writer has taken it chiefly on locust, Ceanothus 
and walnut. 

Subfamily SMILIINtE (Stal). 

This subfamily includes all but the three preceding members of 
our fauna. In all of them the pronotum conceals the scutellum and 
the tibiae are not foliaceous. The seventeen genera occurring in 
Kansas may be separated by the following key: 

KEY TO GENERA. 

A. Tegmina entirely free, not covered by pronotum. 
B. Veins of corium closely united at base. 

C. Suprahumeral horns present. Ceresa. 

CC. Suprahumeral horns absent. Stictocephala. 

BB. Veins of corium widely separated at base. 

C. Tegmina with five apical cells; veins distinct. Acutalis. 

CC. Tegmina with four apical cells; veins indistinct. 

Micrutalis. 
AA. Tegmina partly or entirely covered by pronotum. 

B. Terminal cell of hind wing sessile, its base truncate. 

C. Pronotum without horn or crest. 

D. Dorsum low and rounded. Carynota. 

DD. Dorsum high, compressed, and foliaceous. Archasia. 

CC. Pronotum with horn or crest. 

D. Horn anterior and porrect. Thelia. 

DD. Horn a flat dorsal crest. 

E. Crest arising from between humeral angles. 

Glossonotus. 

EE. Crest arising from behind humeral angles. 

F. Crest strongly step-shaped. Heliria. 

FF. Crest sometimes slightly, but not usually step- 
shaped. 

G. Basal costal cell of tegmina not entirely 
punctate. Telamona. 

GG. Basal costal cell of tegmina entirely 
punctate. Telamoncmthe. 

BB. Terminal cell of hind wing triangular and petiolate. 

C. Base of corium with three veins. 

D. Corium without cross-veins at base. Smilia. 

DD. Corium with cross-veins at base. 

E. Dorsum strongly compressed. Cyrtolobus. 

EE. Dorsum rounded. Ophiderma. 



La\vso.\ : Membracid^ of Kansas. 51 

CC. Base of corium with two veins. 

D. Apical cell of te^;inina transverse. Vanduzea. 

DD. Apical cell of tegmina triangular. 

E. Dorsum strongly elevated, with deep median notch. 

Entylia. 

EE. Dorsum slightly elevated, with weak median de- 
pression. Publilia. 

Genus Ceresa Amyot & Serville. 

To this genus belong our commonest species, most of which are 
green and all of which are at once recognized by their prominent 
suprahumeral horns. 

The seven species known to occur in Kansas may be separated by 
the following key: 

KEY TO SPECIES. 

A. Species brown. 

B. Species larger and very hairy; suprahumerals not recurved; one 
white band on j)osterior process. diceros. 

BB. Species smaller, with few hairs; suprahumerals recurved; two white 
bands on posterior process. albescens. 

AA. Species green. 

B. Dorsal crest marked with brown or reddish. 

C. Suprahumerals short and slightly recur\ed. palmeri. 

CC. Suprahumerals longer and strongly recurved. con-stans. 

BB. Dorsal crest concolorous. 

C. Species small, 7-8 mm. long. 

D. Very hairy species. borecdis. 

DD. Sparsely haired species. brevicornis. 

CC. Species large, 8-10 mm. long. 

D. Suprahumerals long, sloping upward and recurved; cly- 
peus much produced beyond vertex. taurina. 

DD. Suprahumerals stout, nearly straight, clypeus short. 

bubalus. 
Ceresa diceros (Say). 

(PI. Ill, figs. ], 2.) 

Membracis diceros Say, Long's 2nd E.xped., ii, p. 299, 1824; Compl. Writ., i, p. 199. 

Smilia diceros Germar, Silb. Rev. Ent., iii, p. 237, 1835. 

Ceresa postfasciata Amyot & Serville, Hem., p. 540, pi. 10, fig. 3, 1843. 

Ceresa diceros Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, i). 285, 184fi. 

Ceresa vitidalis Buckton, Monog. Membr., p. 172, pi. 36, figs. 3-3b, 1903. 

Funkhouser gives the following technical description: 

Dark brown with trans\erse bands of yellowish white; suprahumeral horns 
stout and blunt; posterior process decurved; tegmina smoky hj'aline. 

Head broader than long, sculptured, basal part strongly and smoothly 
curved, front surface light yellow faintly marked with brown, faintly longi- 
tudinally ridged, very slightly or not at all punctate or pubescent ; eyes 
prominent, extending beyond adjoining lateral margin or pronotum; ocelli 
shining, transparent, nearer to each other than to the eyes; sclerites of front 



52 The University Science Billetin. 

projecting over clypeus at internal angles with a small hook; clypeus strong, 
swollen, roughly three-Iobed, the central lobe the largest, tips strongly hirsute. 

Pronotuni densely and coarsely punctate; anterior surface slightly convex, 
light yellow with numerous brown markings, sparingly pubescent with rather 
long hairs; suprahumeral horns projecting outward and very slightly back- 
ward; lateral surfaces not pubescent, brown with two transverse light bands, 
the anterior broad and irregular in about center, the posterior narrower and 
regular just before apex of j^osterior process; posterior process gradually acute, 
extending beyond internal angles of tegmina. 

Tegmina hyaline, tips sriioky, bases opaque and lightly punctate ; five apical 
and three discoidal cells. Undersurface of body very dark brown. Femora 
dark brown above; tibise and tarsi ferruginous. 

Length, 9 mm.; width between humeral horns, 5.5 mm. 

Internal male genitalia. Styles stout, varying much in the length 
of the cephalic part which, at its greatest length, is shorter than the 
part caudad of the connective, the sharply pointed caudal portion 
bearing two rows of long hairs; connective longitudinally keeled, 
when open nearly twice as long as wide, base concave, apex nar- 
rowed but obtuse; oedagus, viewed laterally, much as in Ceresa 
bubalus, but with dorsal process usually more pointed apically and 
the ventral process distinctly narrower and more acute apically. 

Distribution: Van Duzee reports this species from Nova Scotia, 
Ontario, New York, New Jersey, Pennsylvania, Maryland, North 
Carohna, Ohio, lUinois, Iowa, Kansas, Missouri, Dakota, Colorado, 
Texas, New Mexico and Montana. It is therefore one of our most 
widely distributed species. In Kansas specimens have been taken in 
Douglas, Miami, Neosho, Bourbon, Pottawatomie, Riley, Shawnee 
and Saline counties. 

Hosts: The usual host for this species is elderberry. Funkhouser 
gives the following additional hosts: Locust, oak, sycamore, sweet 
clover, blackberry and butternut. 

Ceresa albescens Van Duzee. 

Ceresa albescens ^'an Duzee, Bui. Buf. Soc. Nat. Sri., ix. p. 3.5, 1908. 
Ceresa bubalus var. a and b Fitch, Homop. N. Y. St. Cab., p. 50, 18.51. 

The following is the original description: 

A little smaller and paler than diceros, to which it is closely allied. Prono- 
tum as in diceros, but with the suprahumeral horns more acute and recurved 
and tipped with black, and the apex longer and more slender. Face, front 
and superior surface of the pronotum greenish- or yellowish-white with scarcely 
a trace of the maculations found in dicero^i; apex of the head less produced, 
the tylus scarcely longer than the cheeks. Sides of the pronotum paler, fer- 
ruginous, becoming somewhat fuscous posteriorly, irrorate with paler and 
marked with a pale marginal line and sometimes with an oblique median 
vitta; protracted apex whitish with a black tip and ferruginous median vitta; 
outer surface of the suprahumerals dark ferruginous, differentiated from the 



Lawsox: AlKMBRACiDiE OF Kansas. 53 

ferruginous sides by the pale lateral line wliich nearly attains the apex of the 
horns. Legs and beneath t'(M-ruginous, the femora darker. Last ventral seg- 
ment of the female a \ tM y little oblique and roimded to the median notch, 
whirli is triangular. acut(\ and reaches the middle of the segment. Plates of 
tlie male not attaming the apex of the anal tube. Elytra almost hyaline, not 
smoky brown as in dircrus, nervures ferruginous. Length, 8-9 mm. 

Distribution. Van Duzee reports this species from Ontario, New 
York, and Kansas. Matauscli took specimens in New Jersey. Tlie 
Kansas specimens were taken at Effingham, Atchison county. 

Hosts. Matausch gives Viburnum as a host. 

Ceresa palmcri Van Duzee. 
(PI. in, fig^. 3, 4.) 

Ceresa palmeri \'aii Duzee, Can. Ent., xl, ii. 114, 1908. 

Funkhouser gives tlie follo\ving technical description: 

Near Ceresa constarus, but differing particularh- in shape of the supra^ 
humeral horns, which are short, terete, and but little recurved; small, reddish 
species, with jironotum rather high, not pubescent. 

Head wider than long, yellowish, only fain>tly sculptured, not punctate; 
eyes prominent, reddish with white borders, extending beyond adjoining lat- 
eral margins of pronotum ; ocelli not prominent, pearly with reddish margins, 
nearer to each other than to the eyes; clypeus continuing lateral margin of 
face, swollen and pubescent at tip. 

Pronotum yellow-green, very strongly marked with brown and reddish; 
dorsal crest curved, strongly marked with red; lateral semicircular impression 
faint, area within it lighter in color than surrounding pronotum; posterior 
process slightly curved downward, about reaching tip of abdomen but not 
extending halfway to extremities of tegmina. 

Tegmina hyaline, wrinkled, bases slightly punctate. L^ndersurface of body 
yellowish. Legs concolorous 3'ellow-green in life, fading to pale yellow in 
cabinet specimens. 

Length, 8 mm.; width. 3.5 mm. 

Internal male genitalia. Styles large and stout, widest just 
caudad of attachment to connective, then broad until suddenly 
narrowed to the acute apex, caudal third with a fringe of long 
hairs extending nearly to the apex on the outer margin, but stop- 
ping considerably cephalad of the apex mesally ; connective elongate 
heart-shaped; cedagus, viewed laterally, with dorsal process of 
medium width, ventral process more heavily chitinized and quite 
stout until near the quite acute apex, the membrane guarding the 
functional orifice large and about midway between the base and 
apex of the process. 

Distribution. Van Duzee reports this species from Ontario, New 
York and North Carolina. Specimens have been taken in Kansas 
in Linn, Bourbon. Hodgeman and Cowley counties. 



54 The University Science Bulletin. 

Hosts. Funkhouser gives young hickory as a host, while Metausch 
took numerous nymphs of the species from Liquidambar. 

Ceresa constans (Walker). 

Thelia constans Walker, List Homop., ii, p. 563, 1851. 
Ceresa constans Stal, Of. Vet. Akad. Forh., xxvi, p. 245, 1869. 
Ceresa suhulata Provancher, Pet. Faune Ent. Can., iii, p. 3.38, 1890. 
Ceresa illinoiensis Goding, Bui. 111. St. Lab. Nat. Hist., iii, p. 404, 1894. 

Funkhouser gives the following technical description: 

Small and distinctly reddL^ih; dorsal cre.st low, median lateral line red; meto- 
pidium convex; horns gharji and much recurved; i)Osterior process nearly 
straight, usually tipped with red; head triangidar; tegmina and wings hyaline. 

Head subtriangular, weakly sculptured, faintly longitudinally furrowed, very 
finely and lightly punctate, not pubescent; eyes prominent, dark brown with 
lighter edges, extending beyond adjoining lateral margins of pronotum; ocelli 
glassy, nearer to each other than to the eyes; clypeus much longer than wide, 
extending for more than half its length beyond lateral margin of face, tip 
hirsute. 

Pronotum deeply and coarsely punctate, not pubescent, median carina 
])rominent; dorsal crest low, rising but little higher than tips of suprahumeral 
horns; horns slender, sharp, hiuch recurved, extending upward and curving 
backward; metopidium convex, regular; lateral semicircular impression deep, 
concolorous; posterior process nearly straight, not reaching the extremity of 
the abdomen and reaching barely one-third the distance to the tips of the 
tegmina. 

Tegmina hyaline. Undersurface of body and legs yellowish. 

Length, 8 mm.; width, 4 mm. 

Distribution. Van Duzee reports this species from Ontario, New 
York, North Carolina and Illinois. Specimens have also been taken 
in Riley county, Kansas. 

Hosts. Funkhouser gives locust as the host. 

Ceresa borealis Fairmaire. 

(PI. Ill, figs. 7, 8.) 
Ceresa borealis Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 284, 1846. 

Funkhouser gives the following technical description: 

Resembling C. bubalus in general outline, but much smaller and very hairy; 
metopidium convex; dorsum curved, posterior process only slightly decurved; 
head impunctate; notch of last ventral segment of female broad and tri- 
angular. 

Head broader than long, yellowish, roughly sculptured, faintly longitudi- 
nally striate, not punctured nor pubescent; eyes prominent, mottled with 
green and brown, extending beyond adjoining lateral margins of pronotum; 
ocelli small, reddish, nuich nearer to each other than to the ej^es; clypeus 
rounded, somewhat protruding, extending for more than half its length below 
lateral margin of face, tip hirsute. 

Pronotum green, finely, deeply and densely punctate, very hairy; meto- 
pidium convex; median carina faint; suprahumeral horns stout, blunt, nearly 



Lawson: Membracid^ of Kansas. 55 

straight, projecting almost dircctlj' outward; dorsal crest regularly arcuate; 
lateral semicircular inii)ression nearly obsolete; posterior process curving 
slightly downward, not extending beyond tip of abdomen and reaching only 
for a short distance beyond internal angles of tegmina. 

Tegmina entirely hyaline, somewhat wrinkled, bases lightly punctate. Legs 
and undersurfaces of bodj' concolorous greenish. 

Length, S mm.; width, 4 mm. 

Internal male genitalia. Style very large when compared with 
the otiier members of the genus, anterior process short, a small but 
distinct knob to the connective, posterior portion extending straight 
until considerably past the connective, then bending rather abruptl}^ 
laterad and ending in a tip that on its mesal margin bears a number 
of small but distinct teeth; connective very long and rather slender, 
notched at base; cedagus, viewed laterally, small, with dorsal proc- 
ess strongly humped, the ventral process with basal knob, then ta- 
pering regularly to acute tip, membrane around functional orifice 
at middle third. 

Distribution. Van Duzee records this species from Ontario, New 
York, New Jersey, Pennsylvania, North Carolina, Ohio, Kansas, 
Colorado, and Utah. In Kansas specimens have been taken in the 
following counties: Cowley, Sumner, Bourbon, Cherokee, Chau- 
tauqua, Douglas, Riley, Shawnee, Crawford, Miami, Linn, Neosho, 
Rawlins, and Ottawa. It is thus seen to extend well over the state. 

Hosts. Funkhouser gives the following host list for this species: 
Wild grape, locust, elder, willow, oak, hickory, pignut, raspberry, 
sycamore, apple, and pear. 

Ceresa brevicornis Fitch. 

Ceresa brevicornis Fitch, Trans. N. Y. St. Agr. Soc, xvi, p. 451, 1856. 

The following is the original description: 

This is so similar to the common Buffalo tree hopper that it will scarcely 
be distinguished from it except by a practiced ej'e, although it is undoubtedly 
a distinct species. It differs from that in having the horns much more short, 
and the sides' of the thorax when viewed in front are not gradually curved out- 
wards, but are straight or rectilinear, with the horns abruptly projecting from 
the comer at the upper end of this line. The acute spine at the tip of the 
thorax is also more long and slender. The thorax between the horns is slightly 
convex. The dried specimen is of a pale dull j-ellow color freckled with faint 
pale green dots and with a paler straw-colored stripe, quite distinct, upon the 
angular sides of the thorax from each eye upward to the horn and from thence 
to the summit of the thorax. 

Length of the female, 0.36. 

It was met with upon hickory bushes in New Jersey. 



56 The University Science Bulletin. 

Distribution. Van Duzee reports this species from Ontario, Con- 
necticut, New York, New Jersey, Pennsylvania, Ohio, and Illinois. 
A single specimen taken at Kansas City, Mo., and determined by 
Dr. E. D. Ball, evidently puts this species within our range. 

Hosts. Van Duzee gives basswood as a host. 

Ceresa taurina Fitch. 

(PI. II, figs. 3, 4.) 

Membracis taurina Harris, List Ins. Mass. in Hitchcock, Geol. Mass., p. 579, 1833 (MS 
name). 

Enchenopa taurina Walker, List Homop., ii, p. 495, 1851 (MS name). 
Ceresa taurina Fitch, Trans. N. Y. St. Agr. Soc, xvi, p. 335, 1856. 

Funkhouser gives the following technical description: 

Slightly smaller than C. bubalus, but resembling it in color; body slender 
and metopidiiim conca\'e transversely; horns sharp, curving upward and back- 
ward. 

Head roughly triangular, wider than long, roughly sculptured, not punctate 
nor pubescent, basal margin strongly curved; eyes prominent, brown and in 
some cases barred with darker, extending beyond the adjoining lateral mar- 
gins of the pronotum; ocelli prominent, pearly, occasionally margined with 
reddish, nearer to each other than to the eyes; clypeus subrectangular, swollen 
and protruding, extending for half its length beyond lateral margin of face, 
faintly trilobed, apex bristled. 

Pronotum deeply and coarsely pimctiu'ed, bright green fading to yellow, 
sparingly pubescent; metopidium strongly concave, with curved, transverse 
margin, area above eyes smooth; suprahvmieral horns slender and sharp, ex- 
tending upward and backward, often much curved, tips generally darker than 
bases; dorsal crest high and strongly curved; semicircular lateral impression 
deep and brownish; posterior process slender, strongly decurved, extending 
beyond apex of abdomen and halfway to tips of tegmina. 

Tegmina and wings entirely hj'aline. Underparts of body and legs yellow- 
green. 

Length, including tegmina, 9 mm.; width between tips of horns, 5.5 mm. 

Internal male genitalia. Styles with anterior part weak, strongly 
narrowed near middle of connective, then much widened into the 
large posterior portion, which ends in an oblique and serrate tip, 
both margins of the caudal half of the posterior portion bearing 
long hairs; connective long and rather narrow, folding longitudi- 
nally, and with the usual basal incision; cedagus, viewed laterally, 
with basal process long and slender, with a distinct knob at point of 
attachment to the connective, the ventral process rather narrow, 
tapering gradually to subacute tip, functional orifice covering the 
middle third of the ventral side. 

Distribution. Van Duzee reports this species from Ontario, Mas- 
sachusetts, New York, New Jersey, Pennsylvania, Virginia, North 



Lawson: Membracid^ of Kansas. 57 

Carolina, Ohio, Michigan, Iowa, Kansas, Colorado, and Arizona. 
Specimens have been taken in Kansas in Sumner, Hodgeman, Doug- 
las, Riley, Shawnee, Ottawa, Linn, Cherokee, Bourbon and Miami 
counties. 

Hosts. Funkhouser reports this species from the following hosts: 
Raspberry, hickory, potato, blackberry, dahlia, hazelnut, locust, 
witch hazel, blue grass, oak, pear, apple, sweet clover, and bitter- 
sweet. Miss Branch adds horseradish and choke cherry. Matausch 
mentions Solidago. 

Ceresa bubalus (Fabricius). 

(PI. II, figs. 1-2 and 5-8.) 

Membracif bubalus Fabricius, Ent. Syst., iv, p. 14, 1794. 
Centrotus bubalus Fabricius, Syst. Rhyng., p. 20, 1803. 
Ceresa bubalus Fitch, Homop. N. Y. St. Cab., p. 50, 1851. 

Funkhouser gives the following technical description. 

Bright green fading to yellowish in cabinet specimens; horns heavy and 
stout, pointing directly outward; metopidium broadly convex; dorsal crest 
high and regularly arched; posterior process slender and recurved; tegmina 
and hind wings entirely hyaline; clypeus heavy, stout, and bristled. 

Head one-third broader than long, longitudinal striate sculpturing; basal 
part broadly curved, front surface yellow, not punctate nor pubescent; eyes 
prominent, dark brown, extending beyond lateral margin of pronotum adjoin- 
ing; ocelli prominent, protruding, with brilliant orange borders, nearer to each 
other than to the eyes; clypeus strong, heavy, continuing lateral outline of 
face, apex bristled. 

Pronotum densely and coarsely punctate; metopidium strongly convex, 
smooth impunctate areas above the eyes, sparingly pubescent with short scat- 
tered hairs; suprahumeral horns stout, blunt, projecting almost directly out- 
ward, not at all upward, tips often brownish, whitish line extending backward 
from tip to lateral margin; lateral surface marked with light-colored semicir- 
cular impression; posterior process slender, depressed, extending half way to 
apices of tegmina and slightly beyond tip of abdomen, apex brownish. 

Tegmina hyahne, bases lightly punctate. Undersurface of body yellowish. 
Legs greenish. 

Length to apices of tegmina, 10 mm.; width between horns, 6 mm. 

Internal male genitalia. Styles seemingly varying considerably 
in size, usually large, the anterior portion about as long as the 
posterior, which ends in a sharp point which is usually perfectly 
smooth, but sometimes bears a few indistinct teeth, the margins 
with two rows of hairs of which the outer one is longer and consists 
of longer hairs; connective about oval in outline, the base incised 
and with a distinct tendency to fold along a longitudinal keel; 
oedagus, viewed lateral!}', wdth the dorsal process varying from 
rather slender and straight to quite stout and hunch-backed, the 



58 The University Science Bulletin. 

ventral process of medium thickness, the apex acute and with the 
usual membrane around the functional orifice near the middle third. 

Distribution. This species occurs throughout the United States 
as shown by the following records of its distribution given by Van 
Duzee: Nova Scotia, Ontario, New Hampshire, Massachusetts, New 
York, New Jersey, Pennsylvania, Maryland, Delaware, North Caro- 
lina, Tennessee, Ohio, Indiana, Illinois, Iowa, Minnesota, Kansas, 
Missouri, Dakota, Montana, Wyoming, Colorado, Utah, New Mex- 
ico, Texas, and California. 

As far as our collecting in this state shows, this species is by far 
the most widely distributed, specimens having been taken in the 
following counties: Douglas, Wilson, Riley, Comanche, Russell, 
Butler, Pawnee, Finney, Graham, Rawlins, Cherokee, Wallace, 
Harper, Shawnee, Wyandotte, Allen, Decatur, Harvey, Saline, Neo- 
sho, Sumner, Reno, Hodgeman, Osborne, Labette, Anderson, Potta- 
watomie, Wabaunsee and Ottawa. 

Hosts. The list of plants on which this species feeds is very 
large, including many weeds and trees. Funkhouser records it 
from the following hosts: Sycamore, aster, poplar, potato, butter- 
nut, hazelnut, pear, sumac, oak, locust, elm, willow, elder, sweet 
clover, hickory, pignut, apple. Miss Branch reports it from osage 
orange, horseradish, gama grass, sunflower and alfalfa. Coding 
records it from apple, potato, tomato, pear, peach, plum, grape, 
apricot, almond, willow, locust and Japanese lily. Gillette and 
Baker report it from willow, apple, soft maple, Solidago spectabilis, 
Aster canescens, Apocynum androscemijolium, alfalfa, and Glycr- 
rhiza lepidota. The writer has taken it most frequently on apple, 
locust, hickory, sweet clover and alfalfa in Kansas. 

. Genus Stictocephala Stal. 

The members of this genus ai^e greenish species, which differ from 
those of the preceding genus by lacking the suprahumeral horns. 

The three species occuring in Kansas may be separated by the 
following key: 

KEY TO SPECIES. 

A. Carinate sides of the metopidium meeting before the middle of the body. 

iiiermis. 
AA. Carinate sides of the metopidium meeting at or behind the middle of 
the body. 
B. Carinate sides of the metopidium meeting at or near middle of 
dorsum. lutea. 

BB. Carinate sides of metopidium meeting much behind middle of 
dorsum. fcstina. 



Lawson: Membracid.e of Kansas. 59 

Stictocephala inermis (Fabriciiis). 

(PI. IV, figs 1, 2, and 5-8.) 

Menibracis inermis Fabricius, Syst. Ent., p. G77, 1775. 

Cicada inermis Gmelin in Linnaeus, Syst. Nat., edn. 13, i, pt. 4, p. 2093, 1788. 
Centrotus inermis Fabricius, Syst. Rhyng., p. 21, 1803. 

Membracis goniphora Say, Jl. Acad. Nat. Sci. Phila., vi, p. 243, 1830; Compl. Writ., 
ii, p. 377. 

Smilia inermis Fitch, Ilomop. N. Y. St. Cab., p. 48, 1851. 

Ceresa gonophora Walker, List Homop., iv, p. 1141, 1851. 

Thelia inermis Walker, List Homop., iv, p. 1142, 1851 (under lutea). 

Stictocephala inermis Stal, Of. Vet. Akad. Forh., xxvi, p. 246, 1869. 

Stictocephala sanguino-apicalis Coding, Bui. 111. St. Lab. Nat. Hist., iii, p. 408, 1894. 

Funkhouser gives the following technical (iescription : 

Fine large species, brilliant green, slowly fading to yellowish in dried 
material; metopidium perpendicular; dorsal crest high and arcuate; posterior 
process slender and curving downward; tegniina and wings entirely hj'aline; 
upper parts of femora often marked with black. 

Head broad, nearly smooth, very finely and faintly punctate, longitudinally 
striate; eyes prominent, subtriangular, very dark bordered with white, ex- 
tending beyond adjoining lateral margins of pronotum; ocelli prominent, 
brownish, nearer to each other than to the eyes; inferior margins of vertex 
broadly sinuate; clypeiis broad, sparingly pubescent, median lobe of apex 
extending below lateral lobes. 

Pronotum densely and coarsely but not deeply punctured ; metopidium con- 
vex, median carina distinct but irregular; sides of metopidium meeting be- 
fore middle of body; lateral semicircular impression deep; posterior process 
long, slender, gradually acuminate, curving downward, extending beyond abdo- 
men and reaching about halfway from internal angles to apices of tegmina. 

Tegmina entirely hyaline, slightly wrinkled, bases greenish and lightly 
punctured. Undersurface of body j'ellowish; segments of abdomen in some 
cases bordered with black; notch of last ventral segment of female broadlj^ 
angular. Femora often marked with black above; tarsi ferruginous. 

Length to tips of tegmina, 9 mm.; width between humeral angles, 4 mm. 

Internal male genitalia. Styles large anci stout, the anterior por- 
tion long and wicie, the posterior part curving to the truncate anci 
serrate tips, which vary from nearly transversely truncate to quite 
obliquely truncate, the apex in the latter case being quite pro- 
nounced, the posterior fourth with two rows of very long hairs, the 
outer row being longer; connective short, widest just behind the 
middle, usually pentagonal in shape; oedagus, viewed laterally, 
much as in Ceresa, the dorsal process varying in width and with a 
small to a very pronounced hump, but with no projection to the 
connective, ventral process moderately stout, slightly concave on 
ventral side preapically, the functional orifice occupying nearly half 
its length medially. 

Distribution. This species is found throughout the United States 



60 The University Science Bulletin. 

as shown by the following distribution giyen by Van Duzee: On- 
tario, New York, New Jersey, Pennsylvania, North Carolina, Ohio, 
Illinois, Iowa, Dakota, Kansas, Missouri, Colorado, New Mexico, 
Arizona, California, and Montana. 

In Kansas it has been taken in the following counties : Chautau- 
qua, Douglas, Hodgeman, Cowley, Ottawa, Riley, Dickinson, Linn, 
Ellis, Bourbon, Montgomery, and Wabaunsee. 

Hosts. Funkhouser records this species from sweet and red 
clover, timothy and apple. Miss Branch mentions gama grass. 
Coding reports it from plum, oats, oak and alfalfa. The writer 
has taken it very commonly on apple. 

Stictocephala lutea (Walker). 

Thelia lutea Walker, List Homop., ii, p. 559, 1851. 

Thelia inermis Walker, List Homop., iv, p. 1142, 1851. 

Gargara pectoralis Emmons, Nat. Hist. N. Y. Ins., p. 157, p. 1, 13, fig. 12, 1854. 

Stictocephala lutea Stal, Of. Vet. Akad. Forh., xxvi, p. 247, 1869. 

Funkhouser gives the following technical description: 

Small species; grass-green above, usually marked with black below; meto- 
pidium sloping, dorsal crest not high, not regularly arcuate; tegmina smoky 
hyaline. 

Head perpendicular, subtriangiilar, broader than long, finely punctate, 
sparingly pubescent, weakly sculptured; eyes prominent, brown usually banded 
with- reddish, extending outward as far as lateral angles; ocelli distinct, yel- 
lowish margined with brown, much nearer to each other than to the eyes; 
inferior margins of vertex weakly sinuate, their ventral raesal angles ending in 
hooks; clypeus robust, extending only slightly beyond inferior margins of 
vertex. 

Pronotum closely and deeply punctate; metopidium convex, median carina 
faint, smooth yellowish area on each side near base of head, sides of meto- 
pidium meeting at or a little before middle of body; dorsal crest not high, 
sloping gradually from junction of carinate edges of metopidium to posterior 
process; semicircular lateral impression weak; posterior process slender, gradu- 
ally acute, extending as far as tip of abdomen and to a point on tegmina half- 
way between internal angles and apices. 

Tegmina hyaline, smoky at apices. Under parts of thorax distinctly black. 
Legs generally marked with black. Notch of last ventral segment of female 
very small or obsolete. 

Length, 6.5 mm.; width, 2 mm. 

Internal male genitalia. Styles stout, especially posterior portion, 
bent in near middle of connective, then flaring widely till just before 
the incurved tips, which are transversely truncate with the inner 
angle prominent and distinctly serrate on both its margins, the 
apical fourth of the styles bearing on each margin a row of long 
hairs; connective large, elongate, widest just caudad of the middle; 
cedagus, viewed laterally, with medium-sized and humped dorsal 



J 



Lawson: Membracid.e of Kansas. 61 

process, the ventral process with a distinct tubercle to connective 
and wide till the end of the functional orifice, then suddenly nar- 
rowed, after which it is nearly parallel-margined to the apex. 

Distribution. Seemingly most abundant in the eastern United 
States as shown by the following records given by Van Duzee: On- 
tario, Connecticut, New York, New Jersey, Pennsylvania, District 
of Columbia, North Carolina, Georgia, Florida and Illinois. There 
are specimens of this species in the Snow collection from Beaver 
Creek, Montana, thus extending its range westward. 

The following Kansas counties have yielded specimens: Linn, 
Montgomery, Neosho, Riley and Douglas. 

Hosts. Matausch reports this species from Solidago; Funkhouser 
from oak and daisy ; Coding from wheat. 

Stictocephala festina (Say). 

(PI. HI, figs. 5, 6.) 

Membracis festina Say, Jl. Acad. Nat. Sci. Phila., vi, p. 243, 1830; Compl. Writ., ii, p. 
377. 

Stictocephala festiva Walker, List Honiop., iv, p. 1141, 1852. 
Stictocephala uniformis Stal, Hemip. Fabr., ii, p. 24, 1809. 
Stictocephala festina Stal, Of. Vet. Akad. Forh., xw-i, p. 246, 1869. 

The following is the original description: 

Thorax with a subacute Hne each side before, meeting behind the middle. 

Inhabits Florida. 

Body yellowish-green; thorax unarmed, carinate behind; at tip attenuated, 
subulate and complying with the general cunature ; each side before a carinate 
line, meeting together at the carina behind the middle, with the carina tinged 
nith rufous; front of the thorax not altogether flat, but a little convex; heme- 
lytra, three temiinal cellules unequal; the two costal ones equal, as broad as 
long; the inner one not Qb\'iousl3- larger than the others together, somewhat 
longer than broad. Length to tip of hemelytra one-fifth of inch. The lateral 
prominent lines of the unarmed thorax, separate this species from all those 
I have described excepting goniphera, which meet before the middle of the 
length of the back. 

Internal male genitalia. Styles quite small, anterior portion 
smaller than posterior, converging posteriorly to the wide posterior 
portions, which, opposite the connective, have a distinct lateral an- 
gle and then narrow but slightly to the obtuse apices, each bearing 
a long outer row and a short mesal row of stout hairs; connective 
quite small and triangular; cedagus, viewed laterally, quite charac- 
teristic, the dorsal process ending in a large rounded lobe which 
bears a small, fingerlike terminal process, the ventral process ending 
in a swollen and obliquely truncate apex which bears the functional 
orifice. 



62 The University Science Bulletin. 

Both the styles, connective and oedagus of this species are so en- 
tirely different from the corresponding parts in the two preceding 
species that it does not seem possible that they could be members 
of the same genus. 

Distribution. This species occurs abundantly in the Southern 
states and in more limited numbers as far north as Connecticut, and 
Canada in the East and Montana in the West. It has been reported 
from Florida, Virginia, Pennsylvania, Georgia, Missouri, Texas, 
Iowa, Montana, Colorado, New York, Connecticut, New Jersey, 
Ottawa, Can., Utah, Arizona, Mississippi, Tennessee, Alabama, 
North Carolina, Louisiana, Kansas, Iowa, Lower California, Mex- 
ico, and the West Indies. 

Hosts. Wildermuth records this species from the following hosts: 
Alfalfa, cowpeas, tomato, almond, Bermuda grass, Johnson grass, 
wheat, barley, oats, bur clover, yellow sweet clover, soy beans, red 
clover, vetch, Hordeum murinum, beans, sunflower, cocklebur, 
Erigeron canadensis, mesquite, cottonwood, Sporobolus airodes and 
Trichlaris mendocina. 

Genus Acutalis Fairmaire. 
In this genus are small species with the prothorax dark and with 
five apical cells in the tegmina, the veins of which are quite distinct. 
A single species is reported from Kansas. 

Acutalis tartafea (Say) 

(PI. V, figs. 5, 6.) 

Membracis tartarea Say, Jl. Acad. Nat. Sci. Phila., vi. p. 242, 1830; Compl. Writ., ii, 
p. 376. 

Ceresa tartarea Walker, List. Homop., iv, p. 1141, 18.52. 

Acutalis tartarea Uhler, Bui. U. S. Geol. Geog. Surv., i, p. 345, 1876. 

Ceresa semicrema Provancher, Pet. Faune Ent. Can., iii, p. 235, 1886. 

Funkhouser gives the following technical description: 

Small elongate species, very black, with eyes, undersurface of body and in 
some cases lateral margins of pronotum white, apices of tegmina abruptly 
hyaline. 

Head twice as broad as long, densely black, smooth, not punctate nor pu- 
bescent; eyes prominent and white; ocelli small, white, about equidistant 
from each other and from the eyes; clypeus foreshortened, smooth, extending 
only slightly in a semicircular curve below inferior line of face. 

Pronotum intensely black above, finely punctate, not pubescent, lateral 
margins and tip of posterior process in some cases marked with white; dorsal 
crest low, weakly convex; posterior process nearly straight, slightly decurved, 
more or less tectiform, extending beyond abdomen and almost to end of apical 
cells of tegmina, but not reaching apex of hyaline border. 

Tegmina opaque black for basal two-thirds, apical third suddenly hyaline; 



Lawson: Membracid^ of Kansas. 63 

veins heav}^ ami black; wide apiral border; bat>al third i)iinctate. Undersur- 
face of body pale. Legs yellowish, tarsi fuscous. 

Length to apices of tegmina, 4.5 mm.; width between humeral angles, 2 mm. 

Internal male genitalia. Styles small, anterior portion slender, 
then wide opposite connective and narrowing again to the rather 
slender apical portions, which are strongly hooked, bear a few hairs, 
and end in an acute apex; connective comparatively large, semi- 
circular; cedagus, viewed laterally, quite large, anterior process 
smaller and ending acutely, posterior process very large and ending 
obtusely. 

Distribution. Van Duzee reports this species from Ontario, 
Massachusetts, New York, Pennsylvania, District of Columbia, 
North Carolina, Florida, Mississippi, Illinois, Iowa, Missouri, Colo- 
rado, and Utah. Miss Branch reports it from Douglas county, 
Kansas. 

Hosts. Miss Branch gives Ambrosia trifida as a host. 

Genus Micrutalis Fowler. 

The members of this genus are small and have but four apical 
cells in the tegmina, the veins of which are very obscure. 
A single member of the genus occurs in Kansas. 

Micrutalis calva (Say). 

(PI. V, figs. 3, 4.) 

Meinbracis calva Say, Jl. Acad. Nat. Sci. Phila., vi. p. 242, 1830; Conipl. Writ., ii, p. 376. 

Membracis melanogranuna Perty, Del. An. Art., pi. 35, fig. 10, 1834. 

Smilia flavipennis Germar, Silb. Rev. Ent., iii, p. 240, 1835. 

Acutalis flavipennis Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 497, 1846. 

Ceresa calva Walker, List Homop., iv, p. 1141, 1852. 

Acutalis melanogramma Walker, List Homop., ii, p. 591, 1851. 

Acutalis calva Fitch, Trans. N. Y. St. Agr. Soc, xvi, p. 391, 1856. 

Acutalis illinoiensis Coding, Can. Ent., xxv, p. 53, 1893. 

Micrutalis illinoiensis Baker, Can. Ent., xxxix, p. 116, 1907. 

Micrutalis calva Baker, Can. Ent., xxxix, p. 116, 1907. 

Funkhouser gives the following technical description: 

Very minute; one of the smallest species of Membracidae in the United 
States; usually strongl^v marked with black, although color is variable; abdo- 
men j-ellowish; tegmina hyaline, veins very indistinct. 

Head broad, smooth, hghtly punctate, not pubescent, upper third black, 
lower two-thirds 3'ellowish; eyes prominent, white or gray; ocelli not promi- 
nent, pearly, about equidistant from each other and from the eyes and situated 
slightly above an imaginary line drawn through centers of eyes; clypeus 
rounded, continuing sinuate outline of inferior margin of face. 

Pronotum low, nearlj^ flat, finelj' punctate, not pubescent, anterior part 
usually black, tip of posterior proce-ss generally pale; posterior process stout, 
triangular, just reaching internal angles of tegmina and not extending as far 
as tip of abdomen. 



64 The University Science Bulletin. 

Tegmina entirely hyaline, not punctate nor pubescent at base, veins indis- 
tinct, apical border broad. Entire abdomen pale; undersurface of thorax of- 
ten marked with black. Femora black or ferruginous; tibiae fuscous, tarsi fer- 
ruginous. 

Length, 3-3.5 mm.; width, 1.5-1.7 mm. 

Internal male genitalia. Styles with anterior and posterior parts 
of about equal length, anterior ends pointed, strongly swollen op- 
posite connective, posterior part curving laterad, with a distinct 
hook before the acute apex, and with about a dozen hairs along 
the lateral margin; connective small, notched basally, widest at the 
truncate apex; oedagus, viewed laterally, U-shaped, the base nearly 
straight and with a large process to connective, functional orifice 
along the side of the posterior arm which bears numerous sawlike 
teeth on its cephalic aspect and a few scattered ones laterally. 

Distribution. This is a very widely distributed species occurring 
throughout the Eastern and Southern states as shown by the fol- 
lowing distribution given by Van Duzee: New Hampshire, Massa- 
chusetts, New York, Pennsylvania, Maryland, District of Columbia, 
Virginia, North Carolina, Georgia, Florida, Alabama, Mississippi, 
Ohio, Michigan, Illinois, Missouri, Kansas, Iowa, Arkansas, Texas, 
Colorado. It is also known to occur in the West Indies. 

Specimens have been taken in the following Kansas counties: 
Reno, Douglas, Riley, Bourbon, Harper, Chautauqua, Cowley, 
Cherokee, Allen, Harvey, Montgomery, Butler, Kingman, Sumner, 
Miami and Ottawa. 

Hosts. Funkhouser gives black locust as a host. It is commonly 
taken when sweeping weeds and grasses in Kansas. 

Genus Carynota Fitch. 

The members of this genus have the elytra partially covered by 
the pronotum, which lacks a horn or crest. The dorsum is low and 
rounded. 

A single species occurs in Kansas. 

Carynota mera (Say). 

Memhracis viera Say, Jl. Acad. Nat. Sci. Phila., vi, p. 301, 1831 ; Compl. Writ., ii, p. 379. 

Carynota mera Fitch, Homop. N. Y. St. Cab., p. 48, 1851. 

Gargara niajus Emmons, Nat. Hist. N. Y. Ins., p. 156, pi. 13, fig. 6, 1854. 

Ophiderma mera Fitch, Trans. N. Y. St. Agr. Soc, xvi, p. 465, 1856. 

Craynota strombergi Goding, Bui. 111. St. Lab. Nat. Hist., iii, p. 443, 1894. 

Funkhouser gives the following technical description: 

Fine large species; gray marked with dark brown and chestnut; pronotum 
convex and elevated; tegmina fuscous-hyaline tipped with dark brown. 

Head nearly twice as broad as long, uniform light gray, very distinctly 
punctate, sparingly pubescent with short, white hairs; eyes very prominent 



Lawson: Membracid^ of Kansas. 65 

and lirowu; ocrlli proiniiunt, pc.-iiiy. iiiai]LiiiuMl wil li i)range. somewhat pro- 
tniding, nearer to each otlier tliaii to the eyes; clypeus subtriangular, con- 
tinuing inferior outline of fac(\ tip i>roduced in small tooth, hirsute. 

Pronotvun gray, finely punctate. ]>ubescent. median carina percurrent ; meto- 
pidium convex, irregular brown mark above internal angle of each eye; dorsal 
line arcuate, suddenly depressed before posterior process in female, depression 
not so evident in male; wide, dark brown, transverse band crossing middle of 
pronotum on each side; jiosterior jirocess heavy, pointed, tip chestnut. 

Tegmina smoky hyaline, veins prominent, bases punctate, especially along 
veins and at costal margins, tips dark brown or black. Legs and undersurface 
of body ferruginous. 

Length: female, 10 mm.; male, 8.5 mm. Width: female, 5 mm.; male 
4 mm. 

Distribution. Van Duzee reports this species from Ontario, New 
York, New Jersey, Pennsylvania, North Carolina, Ohio, Illinois, 
Iowa, Missouri and Texas. The writer took a single specimen in 
Ottawa county, Kansas. 

Hosts. Funkhouser mentions hickory, butternut, pecan and oak 
as hosts of this species. 

Genus Thelia Amyot & Serville. 

The members of this genus are at once recognized by the pos- 
session of a long horn on the anterior part of the prothorax, which 
points upwards and forwards. 

Both members of the genus known to occur in the United States 
are found in Kansas. They may be separated by the following key : 

KEY TO SPECIES. 

A. Prothorax of male without lateral yellow stripe. uhleri. 

AA. Prothorax of male with lateral yellow stripe. bimaculata. 

Thelia uhleri Stal. 

Thelia uhleri .Stal, Of. Vet. Akad. Forh., xxvi, p. 248, 18G9. 

The original description follows: 

Griseo-ferruginea, pilosa, thorace remote pallido-consperso ; tegminibus 
sordide hyalinis, apice fusco-nebulosis, basin versus punctatis. Female: Long. 
9; cum cornu, 13. Lat. 4% mill. Wisconsin. (Mus. Holm.) 

T. himaculatce maxime affinis, pictura thoracis, ejusdem cornu antico 
paullo longiore processuque postico rugis longitudinalibus destituto divergit. 
Caput remote punctulatum. Thorax dense distinctque punctatus, angulis 
lateralibus nonnihil prominulis. rectis, cornu antico antrorsum valde nutante, 
processu postico apicem tegminum subattingent(\ 

Ad hanc speciem verisimiliter spectat Fitch, quiim dicit Membracem belli- 
geram Say ad Theliam referendam esse; hsec species autem ad Platycotem 
pertinet et eadem est ac P. sagittata Germ., quae American! borealem (nee 
Brasiliam) inhabitat. 



66 The University Science Bulletin. 

Distribution. Van Duzee reports this species from Ontario, New 
York, Pennsylvania, Michigan, Illinois, Wisconsin, and Kansas. 

Hosts. Seemingly unknown. 

The writer has not seen specimens of this species, but it is included 
in this paper because of Van Duzee's records. 

Thelia bimaculata (Fabricius). 

(PI. V, figs. 1, 2.) 

Membracis bimaculata Fabricius, Ent. Syst., iv, p. 10, 1794. 
Hemiptycha blnotata Harris, Kept. Ins. Mass., p. 179, 1841. 
Hemiptycha acuviinata Harris, Rept. Ins. Mass., p. 179, 1841. 
Thelia biinaculata Amyot & Serville, Hemip., p. 541, 1843. 
Thelia unanimis Walker, List Homop., ii, p. 566, 1851. 

Funkhouser gives the following technical description: 

Female. Gray with indistinct darker irregular markings; porrect cylindrical 
horn slightly flattened and somewhat darker in color at tip; tegmina hyaline, 
apices fuscous, almost reaching extremity of dorsal process. 

Head, including eyes, twice as broad as long, grayish-yellow mottled with 
ferruginous and brown; margins of lorae strongly sinuate; eyes dark brown; 
ocelli white, nearer to each other than to the eyes and situated on a line drawn 
through centers of eyes; clypeus pilose; beak extending to posterior coxae; 
head very sparingly punctate and sparsely pilose. 

Thorax gray, deeply and densely punctate; median percurrent brown line 
sharpened into a ridge on extremity of horn and at apex of posterior process; 
sides of prothorax roughly and irregularly carinate; horn porrect and greatly 
variable in length; cylindrical except at extreme tip, where it is flattened 
laterally; posterior process heavy, tectiform, gradually acute, almost straight, 
very slightly decurved and extending beyond apices of tegmina. 

Tegmina hyaline, apices fuscous, bases and costal regions lightly' punctate; 
underwings hyaline, two-thirds as long as tegmina. Undersurface of body 
gray-brown, pubescent. Legs uniform yellow-brown; femora thick and 
smooth; tibiae and tarsi densely pilose. 

Length, 11 mm., including horn, 14 mm.; width between humeral angles, 
5.5 mm. 

Male: Differs from female in size and' markings. Smaller, body some- 
what less robust; porrect horn usually shorter and tending to curve; tegmina 
equalling apex of posterior process. Color deep chocolate brown; porrect 
horn almost black; apex of posterior process becoming cinnamon brown; a 
wide, brilliant, lemon-yellow longitudinal stripe on each side of prothorax, 
extending from margin halfway to median dorsal line, also small patches of 
yellow on metopidium; head yellow with brown patches. Undersurface of 
abdomen darker than in female. 

Internal male genitalia. Styles with anterior portion short, 
posterior part longer, wide, and parallel-margined till near apices, 
which are curved strongly laterad and end truncately but with a 
distinct recurved hook, the curved apices bearing scattered short 
hairs; connective large, rather distinctly seven-sided; oedagus, viewed 
laterally, with a slender process to anal tube, the apical portion 



Lawson: Mkmbracid.e of Kansas. 67 

very large, club-f^haped, tlie runctional orilice just before the ex- 
treme apex, the anterior aspeet of the apex with many filelike teeth. 

Distribution. Van Duzee reports this species from Ontario, 
Massachusetts, New York, New Jersey, Pennsylvania, North Car- 
olina, Ohio and Illinois. The writer has taken it in Douglas county, 
Kansas. 

Hosts. Black locust seems to be the only host. 

Genus Glossonotus Butler. 
The members of this genus possess a tongue-shaped crest, which 
arises from between the humeral angles. 

A single species of the genus is recorded from Kansas. 

Glossonotus acuminatus (Fabricius). 

Membracis acuminata Fabricius, Syst. Ent., P. 675, 1775. 

Cicada acuminata Ginelin in Linnaeus, Syst. Nat., edn. 13. i, pt. 4, 2094, 1778. 

Centrotus acuminata Fabricius, Syst. Rliyng., p. 18, 1803. 

Thelia acuminata Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 310, pi. 5, fig. 15, 1846. 

Hemiptycha acuminata Harris, Treat. Ins. Inj. A'eg., edn. 3, p. 223, 1862. 

Telamona acuminatus Stal, Hemip. Fabr., ii, p. 115, 1869. 

Glossonotus acuminata Butler, Cist. Ent., ii, p. 222, 1877. 

Thelia crataegi Smith, Ins. N. J., p. 441, 1890. 

Funkliouser gives the following technical description: 

Dark gray mottled with brown; dorsal crest high, flattened and swollen at 
tip; humeral angles prominent and triangular; tegmina hyaline tipped with 
brown, veins punctured. 

Head almost as long as wide, gray with distinct scattered black punctures 
and fine whitish pubescence; base sinuate; eyes large, prominent, brown, 
extending as far as bases of humeral angles; ocelli large, prominent, pearly 
with white margins, nearer to each other than to the eyes; clypeus continuing 
inferior line of face, punctate with black, pubescent with white, tip prolonged 
into a point; antennae long and well developed. 

Pronotum dark gray with irregular markings of brown, coarsely and regu- 
larh' punctate with black, very sparingly pubescent; metopidium convex, 
median carina prominent and decorated with alternate lines of brown and 
yellowish, irregular black markings above internal angles of eyes, humeral 
angles prominent, triangular, flattened, acute; pronotal crest almost as high 
as length of pronotum, widened and flattened at tip, margin decorated with 
pale areas,- projecting usually forward as well as upward; posterior process 
gradually acuminate, reaching apices of tegmina. 

Tegmina hyaline, tips clouded with smoky brown, bases and margins of 
veins punctate, veins prominent. Undersurface of thorax fuscous; abdomen 
ferruginous. Legs fuscous marked with brown. 

Length. 10 mm.; width between tips of humeral angles, 6 mm. 

Distribution. Van Duzee reports this species from Ontario, Mas- 
sachusetts, New York, New Jersey, Pennsylvania, Michigan, Iowa, 
Kansas and Arkansas. 

Hosts. Funkhouser gives young white oak as the host. 



68 The University Science Bulletin. 

Genus Heliria Stal. 

The members of this genus are characterized by having a dis- 
tinctly step-shaped dorsal crest. 

Two members of the genus are known to occur in Kansas. These 
Van Duzee separates by the following key : 

KEY TO SPECIES. 

A. Larger, 12-13 mm.; grey or brownish grey; posterior foliole of the crest 
Httle elevated above the dorsal line anteriorly, its hind angle subacute; 
humeral angles greatly produced. cristata. 

AA. Smaller, 8 mm., darker brown; posterior foliole of the crest, when well 
differentiated, elevated at least its own width above the dorsal line. 

scalaris. 
Heliria cristata (Fairmaire). 

Thelia cristata Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 311, pi. 5, fig. 14, 184fi. 
Telavxona jagi Fitch, Homop. N. Y. St. Cab., p. 51, 1851. 

Telamona acclivata Emmons, Nat. Hist. N. Y. Ins., p. 155, pi. 3, fig. 5, 1854. 
Heliria cristata Stal, Of. Vet. Akad. Forh., xxiv, p. 556, 1867; .xxvi, p. 249, 1869. 
Telamona cristata Fowler, Biol. Centr. Am., Homop., ii, p. 144, pi. 9, fig. 6, 1896. 

The following is the original description : 

T. cristata. Mexique. Long. 0.012. 

Praecedente differt testaceo-obscuro et tuberculorem dispositione ; primo 
antice inclinato, secundo sat fortiter acuto. 

Xe differe de I'espece precedente que par se couleur d'un testace obscue, 
et la disposition des lobes dorsaux dont I'anterieur est incline en avant, et le 
second plus aigu. Coll. Signoret. 

The preceding species referred to above is H. scalaris. 

Distribution. Van Duzee reports this species from New York, 
New Jersey, North Carolina, and Illinois. Popenoe reports it from 
Kansas. 

Hosts. Unknown. 

Heliria scalaris (Fairmaire). 

Thelia scalaris Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 311, 1846. 

Telamona jagi Fitch, Homop. N. Y. St. Cab., p. 51, 1851. 

Thelia scalaris Walker, List Homop., ii, p. 565, 1851. 

Heliria scalaris Stal, Of. Vet. Akad. Forh., xxiv, p. 556, 1867; xxvi, p. 249, 1869. 

Telamona scalaris Butler, Cist. Ent., ii, p. 222, 1877. 

Funkhouser gives the following technical description: 

A small species, uniform brown in color; crest as high as its length at base;' 
posterior process not reaching apices of tegmina; tegmina smoky hyaline, tips 
browTi. 

Head as wide as long, sculptiu-ed. yellowish, irregularly punctuate with 
brown, sparingly pubescent; base strong^ sinuate; eyes prominent, brown, 
reaching base of humeral angles; ocelli prominent, translucent, nearer to each 
other than to the eyes; clypeus extending below inferior margin of face, 
j'ellowish, punctin-ed with brown, pubescent. 

Pronotum imiform brown, coarsely punctured; dorsal crest swollen at base, 
flattened at apex, as high as its length at base, distinctly step-shaped, anterior 



Lawson: Membracid.e of Kansas. 69 

lobe rouiultHl ami inDjei'tin^ I'orwanl, i)Ot>tciior lobe sharply angular, two-thirds 
as hifih as anterior, both lobes in some cases margined with patches of darker 
brown; posterior process short, heavy, acute, not reaching apices of tegmina; 
humeral angles triangular, flattened, blunt. 

Tegmina smoky hyaline, bases dark brown and punctate, tijis brown, veins 
heavy and often ])unctured along margins, l^ndersurface of thorax ferruginous, 
segments margined with paler; abdomen brown. Legs ferruginous; tibiae and 
tai-si hairy. 

Length, 8 mm.; width, 4.8 mm. 

Distribution. Van Diizec reports this species from Ontario, New 
York, Ne\v Jersey, Pennsylvania, Illinois and Colorado. There is a 
specimen in the Snow collection from Kansas City, Mo., so it un- 
doubtedly occurs in eastern Kansas. 

Hosts. Unknown. 

Genus Telamona Fitch. 

The following is the original description of the genus: 

Humeral angles projecting, pointed and earlike: dorsum compressed-folia 
ceous, the keel abruptly elevated at one or both its ends, forming a somewhat 
square crest or foliole; thorax nearly or quite reaching the tips of the elytra, 
with elevated longitudinal lines on each side; apical cellule triangular, its end 
rounded. The squarish dorsal crest forms a marked distinction between the 
genus here proposed, and that of Thelia, to which it is mo&l nearly related. 

Eleven members of this genus are known to occur in Kansas. 
These may be separated by the following key : 

KEY TO SPECIES. 

A. Anterior margin of crest sloping to front of metopidium, without obvioua 
sinus at its anterior base. 
B. Crest, if elevated, more or less rounded or pointed at apex. 
C. Crest distinctly elevated. 

D. Brownish species; crest higher and triangular. 

pyramidata. 
DD. Greenish species; crest an obtusely conical hump. 

viridia. 
CC. Crest scarcely elevated, broadly rounded above. obsoleta. 
BB. Crest elevated, square or nearly so at apex. 
C. Crest not as high as broad. 

D. Greenish-brown species. lugubris. 

DD. Gray, with oblique brown fascia. decorata. 

CC. Crest as high or higher than broad. querci. 

A A. Crest nearly vertical before or sometimes overhanging. 
B. Color greenish or j^ellowish. 

C. Females bright green; males yellow, banded with brown. 

unicolor. 
CC. Both sexes of about the same color. 

D. Pronotal hump very high. extrema. 

DD. Pronotal hump moderate. collina. 



70 The University Science Bulletin. 

BB. Color not green. 

C. Yellow, mottled with brown. tristis. 

CC. Gray, with transverse brown band. ampelopsidis. 

Telamona pyramidata Uhler. 

(PI. V, figs. 7, 8.) 

Telamona pyramidata Uhler, Wheeler's Rept. Chief Eng. for 1877, p. 1333. 
Telonaca pyramidata Ball, Proc. Biol. See. Wash, xxxi, p. 28, 1918. 

Funkhouser gives the following technical description: 

Long, narrow body; crest triangular and pyramidal, as the name would 
suggest; mottled brown with a dark transverse fascia extending from tip of 
crest to lateral margin of pronotum, and a second shorter fascia behind it; 
posterior process extending to tips of tegmina; tegmina hyaline, punctate at 
bases, brown at apices. Differs from T. declivata chiefly in shape of dorsal 
crest. 

Head wider than long, yellowish with large irregular punctures of brown, 
sparingly pubescent; base regularly sinuate; eyes large, prominent, gray; ocelli 
large, prominent, somewhat protruding, translucent; clypeus subtriangular, 
sutures distinct, apex slightly produced, hairy. 

Pronotum deeply punctate, not pubescent; metopidium convex, decorated 
with patches of yellowish and dark brown, median carina prominent, heavy, 
black, broken by circular areas of yellowish ; humeral angles prominent, tecti- 
form, blunt, brownish; dorsal crest triangular, rounded at tip, margin flattened 
and brown, posterior margin pale ; posterior process long, slender, slightly curv- 
ing downward, extending beyond tips of tegmina; median carina percurrent. 

Tegmina hyaline, bases and costal margins coarsely punctate but not pu- 
bescent, tips brown. Undersurface of thorax flavous; abdomen dark brown. 
Legs yellowish ; tibiae mottled with brown, hairy ; tarsi flavous ; claws fer- 
ruginous. 

Length, 9-11 mm.; width, 5-6 mm. 

Internal male genitalia. Styles with anterior portion broad and 
flat, posteriorly curved outward and with a large, blunt apical hook, 
the apical fourth with a few short hairs ; connective large and heart- 
shaped, bearing three longitudinal ridges; oedagus, viewed laterally, 
U-shaped, anterior process rather slender and with a distinct tooth 
on its cephalic aspect, posterior process large and club-shaped, the 
apex with sawlike teeth on its anterior surface, and the functional 
orifice on the posterior. 

Distribution. Gillette and Baker report this species from Colo- 
rado and Funkhouser from New York. The writer has taken it at 
Ames, Iowa. Specimens have been taken in Kansas from Johnson, 
Pawnee, Cherokee, Riley and Pottawatomie counties. It seemingly 
occurs throughout the Eastern states and west to the Rocky Moun- 
tains. 

Hosts. Funkhouser reports it on chestnut oak. Gillett and Baker 



Lawson: Membracid.e of Kansas. 71 

give Cottonwood and Virginia creeper. The writer has taken it 
abundantly on willow. 

Tlie writer has been unable to find any characters that would 
seem to separate this species from the members of the genus Tela- 
mona. The genitalia are precisely as in other members of the 
genus. Hence it is our belief that the genus Telonaca should be 
sunk as a synonym of Telamona. 

Telamona viridia Ball. 

(PI. VII, figs. 7, 8.) 
Telamona viridia Ball, Proc. Biol. Soc. Wash., xvi, p. 178, pi. 1, fig. 3, 1903. 

The original description follows: 

Resembling pyramidata in size and form, but with less of a hump. Grass- 
green, the male with some fuscous on posterior half of hump and again at 
apex of pronotum. 

Length: female, 11 mm.; male, 9 mm. Width, female, 5.3 mm. 

Pronotal hump in the shape of an obtuse pyramid one-third the distance 
back from eye to apex of pronotum, a slight angle on posterior margin just 
below apex especially marked in the male. Height of hump slightly less than 
one-third the pronotal length. Humeral angles broad, slightly rounded, a 
trifle longer than eye. 

Color. Female, grass-green slightly mottled with yellow; carina light ex- 
cept at apex of hump and at tip, where it is tawny. Male, grass-green; carina 
light interrupted with tawny; a fuscous band nans obliquely backward from 
apex of hump and fades out before reaching the pronotum proper, or some- 
times connects with a tawny spot on lower margin, whole apex of pronotum 
tawny. 

Internal male genitalia. Styles large, anterior end flattened, pos- 
terior part stout, curved strongly laterad apically, bluntly hooked, 
and bearing a few scattered hairs; connective large, heart-shaped, 
with three longitudinal ridges; oedagus, view^ed laterally, U-shaped, 
anterior arm more lightly chitinized and with short blunt anterior 
tooth, posterior arm club-shaped, the apex anteriorly with many 
small filelike teeth, posteriorly with the functional orifice. 

Distribution. Ball reports the type specimens from Colorado, 
Iowa and Illinois. There is a specimen in the Snow collection, taken 
by F. X. Williams, from Gove county, and a single specimen has 
been taken in Riley county, Kansas. 

Hosts. Ball gives cottonwood {Populus monilifera) as the host. 

Telamona obsoleta Ball. 

Telamona obsoleta Ball, Proc. Biol. Soc. Wash., xvi, p. 178, pi. 1, fig. 2, 1903. 

The following is the original description: 

Resembling irrorata, but smaller and with a smaller and more rounding 
hump. Length: female, 10 mm.; male, 9 mm. Width, 5 mm. 



72 The University Science Bulletin. 

Dorsal hump low and much inflated; it scarcely narrows from the base to 
just before the apex where it rounds in to form a carina. Anterior margin 
rising just back of the humeral angles and extending from there half way to 
the apex of the pronotum. The height is about equal to the whole length 
and it rounds down to the pronotum proper at both extremities. Front much 
elevated above the level of the eyes so that the ocelli are farther from the 
base of front than from each other. 

Color. Yellow with the punctures fuscous, sometimes coalescing into brown- 
ish fuscous spots, giving the whole insect an irrorate and mottled appearance, 
with little regularity of pattern. Usually there is a semicircle of lighter shade 
back of the humeral angles and a light spot on middle of hump. There is a 
pair of large, straggling black marks above and within the eyes, some brown 
on the inner nervures of corium, and a smoky brown cloud at apex. 

Distribution. Ball reports this species from Iowa and from Kan- 
sas. Van Duzee records it from Illinois and Fmikhouser from New 
York. In Kansas it has been taken in Pottawatomie and Mont- 
gomery counties. 

Hosts. Ball gives elm as a host; Funkhouser reports it from oak. 

Telamona lugubris Ball. 

Telamona lugubris Ball, Proc. Biol. Soc. Wash., xvi, p. 179, 1903. 

The original description follows: 

Form of reclivata nearly, slightly shorter and stouter built and with a lower 
and longer hump and lacking the markings of that species. Obscurely greenish 
brown. Length: female, 11 mm.; male, 9.5 mm. Width, 5.5 mm. 

Dorsal hump of moderate size, arising just back of lateral angles; anterior 
margin sloping back, forming a right angle with the inclined crest, posterior 
margin perpendicular or slightly overhanging. Base of hump occupying a 
little over two-fifths of distance from humeral angles to apex of pronotum. 
Humeral angles blunt and obtuse, about two-thirds as long as the eye. 

Color. Pale yellow, the more or less darkened punctures giving the insect 
a general grayish cast with still darker shadings on the lateral faces of the 
hump and sometimes on the apex of pronotum. 

Distribution. Ball reports this species from Iowa and also from 
Pottawatomie county, Kansas. Doctor Funkhouser has specimens 
from New Mexico. 

Hosts. Ball gives scrub oak as the host. 

Telamona decorata Ball. 

Telamona decorata Ball, Proc. Biol. Soc. Wash., xvi, p. 179, 1903. 

The original description follows: 

Smaller than lugubris, with a shorter and more rounding hump. Yellowish 
fuscous with the hump deep testaceous brown. Length, 9 mm.; width, 4.5 mm. 

Dorsal hump sloping uj) from both front and rear, crest rounding, highest 
just in front of the middle, hump occupying scarcely two-fifths of the pronotum 
from the humeral angles back. Humeral angles short and blunt, about tw)- 
thirds the length of the eye. 



Lawsox: Membracid^ of Kansas. 73 

Color. Face and pronotiini pale yellow, very slightly washed with brown 
in the female, and with a definite brown shade in the male; median carina 
alternately light and dark l^efore the iiuni)). Hiimii rieli testaceous with a few 
light spots on the sides, a definite light mark at the base in front, which may 
extend up on to the carina, and the whole posterior margin light. This latter 
light spot extends down on to the pronotum and connects with an irregular 
transverse light band about halfway to apex. Each side of this band is an 
irregular testaceous band, the anterior one connected with the testaceous 
hump. A pair of spots above the eyes and the apex of elytra brownish fuscous. 

Distnbution. Ball described this species from specimens taken 
in Iowa, Arkansas, and Pottawatomie county, Kansas. Fimkhouser 
reports it from New York. 

Hosts. Ball mentions red oak as a host. 

Telamona querci Fitch. 

Telamona querci Fitch, Homop. N. Y. St. Cab., p. 51, 1851. 

Telamona quercus Walker, List Homop., iv, p. 1145, 1852. 

TheUa quercus Smith, Cat. Ins. N. J., edn. 1, p. 441, 1891. 

Telamona brunneipcntiis Buckton, Monog. Membr., p. 197, pi. 43, fig. 1, 1903. 

Funkhouscr gives the following technical description: 

Very close to T. monticola; pronotum shorter, darker; dorsal crest with 
prominent pale fascia on posterior margin; tegmina nearh' hyaline, tips 
faintly clouded. 

Head roughly sculptured, flavous mottled with brown, faintly longitudinally 
striate, ^■erJ' faintly punctate, pubescent, base weakly sinuate; eyes prominent, 
dark bro^Mi; ocelli very prominent, protruding, brownish, margins pale, much 
nearer to each other than to the eyes ; clypeus nearly flat, punctate, pubescent, 
base marked with brown, tip extended below inferior margin of face. 

Pronotum densely but finely punctate, sparingly pubescent, dark brown 
mottled with green; metopidium convex, median carina prominent, black in- 
terrupted with pale green ; humeral angles short and blunt ; dorsal crest sloping 
backward, longer than high, higher before than behind, posterior margin dis- 
tinctly pale; posterior process short, acute, marked with greenish before apex, 
not reaching tips of tegmina. 

Tegmina hyaline, bases punctured but not pubescent, tips clouded with 
brown, veins brown. Undersurface of body brown. Legs flavous; tibiae haiiy. 

Length of pronotum, 9 mm.; to tips of tegmina, 11 mm.; width, 5.5 mm. 

Distribution. Van Duzee reports this species from Ontario, New 
York. District of Columbia, Ohio, Colorado, New Mexico, and Ne- 
vada. Popenoe reports this species from Kansas, and there is a 
specimen in the Snow collection from Kansas City, Mo., so that it 
doubtless occurs in the state. The writer took a single specimen of 
the species at St. Paul, Minn. 

Hosts. Funkhouser gives white and chestnut oak as hosts. 



74 The University Science Bulletin. 

Telamona unicolor Fitch. 

Telamona unicolor Fitch, Homop. N. Y. St. Cab., p. 50, 1851. 
Telamona fasciata Fitch, Homop. N. Y. St. Cab., p. 50, 1851. 
Hemiptycha diffusa Walker, List Homop., Suppl., p. 143, 1858. 

Funkhouser gives the following technical description: 

Females large, brilliant iirufonn grass-green; males smaller, bright yellow 
with deep brown fascia. Very striking in color; large size; crest high and 
square; tegmina tipped with brown. 

Female. Head nearly twice as wide as long, green punctate with brown, 
finely pubescent; eyes large, brown; ocelli large, orange, nearer to each other 
than to the eyes; clypeus deeply punctate, pubescent, tip in a pointed ex- 
tension. 

Pronotum concolorous green, fading to mottled yellow in cabinet specimens ; 
very finely punctate and pubescent; metopidium more or less angular, median 
carina distinct, three small brown spots mesad of humeral angles; humeral 
angles produced, triangular, blunt; crest large, high, much higher before than 
behind, anterior margin less sloping than posterior, dorsal margin brownish; 
posterior process long, gradually acute, apex brownish and not reaching tips 
of tegmina. 

Tegmina brownish hyaline, bases and costal regions punctate with black, 
tips clouded with dark brown, veins prominent. Undersurface of thorax 
flavous, abdomen yellowish, pubescent, ovipositor brown. Legs flavous; tibiae 
mottled with brown; tarsi ferruginous. 

Length, 11 mm.; width, 6 mm. 

Male. Differs from female in size and color. Head mottled brown and yel- 
low, much darker than that of female, much sculptured, inferior line of face 
strongly sinuate. 

Pronotum bright yellow, metopidium strongly shaded with brown; dark 
brown fascia on front of dorsal crest; dark brown fascia on posterior third of 
crest extending gradually narrowed to lateral margin of pronotum; posterior 
median line of crest yellow, transverse band of yellow behind crest; apex of 
posterior process brown. 

LTndersurface of body deep brown. Legs strongly fiavous marked with 
brown. 

Length, 10 mm.; width, 5 mm. 

Distribution. Van Duzee reports this species from Ontario, New 
York, New Jersey, Pennsylvania, North Carolina, Michigan, Il- 
linois, Iowa, Kansas, Missouri and Texas. In Kansas it has been 
taken in Pottawatomie county. 

Hosts. Funkhouser gives hickory, butternut, walnut and bass- 
wood as hosts. 

Telamona extrema Ball. 

Telamona extrema Ball, Proc. Biol. Soc. Wash., xvi, p. 179 pi. 1, fig. 1, 1903. 

The original description follows : 

Form of unicolor nearly, smaller, and with a still longer hump. Greenish 
testaceous. Length: female, 10 mm.; male, 9 mm. Width, 5 mm. 



Lawson: Membracid.e of Kansas. 75 

Pronotal hump vpry high, almost quadrate, occupying the anterior three- 
fifths of pronotum, anterior margin rising perpendicularly from face, crest 
highest just back of the well rounded anterior angle from which it slopes 
slightly to the almost perpendicular posterior face. Humeral angles moderate, 
as long as the eyes. 

Color: Greenish testaceous; a spot above each eye and the median carina 
back to the posterior angle of hump fuscous; posterior face of hump broadly 
marked with creamy white, which narrows to a hne on the carina posteriorly 
in the female, and disappeai-s entirely in the male. The lower margin of the 
humeral angles is sometimes marked with fuscous. 

Distribution. This species was described from specimens taken in 
Iowa, and Marion comity, Kansas. Van Duzee reports it also from 
Massachusetts, Rhode Island and New Jersey. 

Hosts. Ball gives oak as a host. 

Telamona collina (Walker). 

Thelia collina Walker, List Homop., ii, p. 565, 1851. 

Telamona collina Butler, Cist. Ent., ii, p. 220, 1877. 

Telamona turritella Buckton, Monog. Membr., p. 198, pi. 44, fig. 6, 1903. 

The following is the original description: 

Testacea vel viridi-flava; prothorax ai)ice niger; carina ferruginea vel fulva; 
alse limpidse; alae anticte basi fulv». 

Testaceous, shining: head finely punctured, short, almost transversely 
spindle-shaped, much narrower than the fore-chest, impressed on each side of 
the disk, with five slight undulations along the hind border, and seven on the 
fore border; a slight fuiTow extends from the hind border to the face, whose 
hind side is slightly obconical and occupies less than half the length of the 
face: fore-chest roughly punctured, convex and with a slight middle ridge 
in front, forming on each shoulder a conical, flat, very prominent horn; keel 
veiy deep behind the shoulders, conical and slightly inclined forward, veined 
along the lower side, slightly undulating and declining abruptly along half 
its length, straight and slightly attenuated from thence to the tip, which is 
black and extends far beyond the tip of the abdomen; ridge mostly ferrugin- 
ous; sides slightly tumid; wings colorless; veins tawny; fore-wings partly 
brown along the hind borders and at the tips, tawny and punctured towards 
the base. Length of the body, 4 lines; of the wings, 10 lines. 

New York. 

Var. /3. Head and fore part of the fore-chest pale yellow tinged with green ; 
keel ta\\Tiy, partly green; legs tinged with green; oviduct pitchy, curved. 

St. John's Bluff, E. Florida. 

Distribution. Van Duzee reports this species from New York, 
Pennsylvania and Florida. It has also been taken in Johnson 
county, Kansas. 

Hosts. Mrs. Slosson and Van Duzee report it from sycamore. 



76 The University Science Bulletin. 

Telamona tristis Fitch. 

Telamona tristis Fitch, Homop. N. Y. St. Cab., p. 51, 1851. 

Telamona coryli Fitch, Honiop. N. Y. St. Cab., p. 51, 1851. 

Telamona spreta Coding, Bui. 111. St. Lab. Nat. Hist., iii, p. 417, 1894. 

Funkhouser gives the following technical description: 

Near T. ampelopsidis in appearance, but smaller and lighter and differing in 
coloration; crest high and square, higher before than behind; tegmina hyaline 
tipped with brown; pronotum yellow mottled with red-brown. 

Head subquadrate, yellowish, faintly longitudinally striate, finely punctate, 
closely pubescent, faintly mottled with brown; eyes prominent, brown; ocelli 
pearly, nearer to each other than to the eyes; clypeus pubescent, tip slightly 
extending below inferior margin of face. 

Bronotum densely punctate, not pubescent, ground color light yellow, a 
broad transverse reddish-brown fascia nearly covering metopidium, a second 
on front of crest, and a third extending down posterior third of crest and reach- 
ing lateral margin of pronotum ; humeral angles produced, triangular, flattened, 
blunt, tips dark; dorsal crest nearly square, truncate at tip, posterior margin 
pale; posterior process long, sharp, not quite reaching tips of tegmina. 

Tegmina smoky hyaline, bases opaque and punctate, tips brown. Under- 
surface of thorax flavous; abdomen brown. Legs ferruginous. 

Length, 8.5 mm.; width, 5 mm. 

Distribution. Reported from Ontario, New York, New Jersey, 
Pennsylvania and Illinois. There are specimens in the Snow collec- 
tion from Kansas City, Mo., so it doubtless occurs in Kansas. 

Hosts. Funkhouser reports this species from hazelnut and oak. 

Telamona ampelopsidis (Harris). 

Membracis cissi (Harris MS), List Ins. Mass., in Hitchcock, Geol. Mass., p. 584, 1833. 
Membracis ampelopsidis Harris, Rept. Ins. Mass., p. 181, 1841. 
Thelia cyrtops Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 310, 1846. 
Telamona ampelopsidis Fitch, Homop. N. Y. St. Cab., p. 51, 1851. 
Telamona monticola Uhler, Stand. Nat. Hist., ii, p. 225, fig. 302, 1884. 

Funkhouser gives the following technical description: 

Fine, large, well-marked species; crest high, erect, front margin nearly per- 
pendicular, hind margin sloping; ground color grayish with brown transverse 
fascia across metopidium, deep brown area at frontal base, brown fascia extend- 
ing from posterior tip of crest to lateral margin of pronotum; tegmina hyaline, 
with brown tips. 

Head yellowish, faintly marked with brown below, sculptured, finely punc- 
tate, sparingly pubescent; eyes prominent, grayish brown; ocelli large, yellow- 
ish, nearer to each other than to the eyes; clypeus smooth, pubescent, tip tri- 
angular. 

Pronotum finely punctate, very sparingly pubescent; metopidium yellow at 
frontal margin, black spot above each eye, median carina prominent, black; 
humeral angles prominent, blunt, extending beyond the eyes as far as the 
length of the eyes; dorsal crest higher before than behind, margin somewhat 
flattened; posterior process long, strong, heavy, extending almost to tips of 
tegmina. 



Lawson: Membracid.e of Kansas. 77 

Teginina hyaline, lightly punctate at base and along costal margins, tips 
brown. Undersurface of body generally uniform gray-brown. 

Male smaller and darker than female, often without characteristic markings. 

Length: female, 10 mm.; male, 8-9 mm. Width: female, 6 mm.; male, 
5 mm. 

Distribution. This species has been reported from Massachusetts, 
New York, New Jersey, Maryland, North Carolina, Illinois, Kansas 
and Colorado. It has been taken in Kansas in Riley, Leavenworth 
and Douglas counties. 

Hosts. Occurs only on Psedera quinquefolia, our common Virigina 
creeper. 

Genus Telamonanthe Baker. 

The members of this genus greatly resemble those of the preced- 
ing genus in general appearance, but they are smaller insects as a 
rule and are all characterized by having the basal costal cell of the 
tegmina completely punctate. 

One species of the genus has been taken in Kansas and another at 
Kansas City, Mo., so that both doubtless occur in the state. 

These species may be separated by the following key: 

KEY TO SPECIES. 

A. Crest nearly quadrate, humeral angles very long. rileyi. 

A A. Crest rounded, humeral angles short. modesta. 

Telamonanthe rileyi (Coding). 

Telamona rileyi Coding, Ent. News, iii, p. 108, 1892. 

Telamona coqnilletti Coding, Bui. 111. St. Lab. Nat. Hist., iii, p. 420, 1894. 

Telamotmnlhe rileyi Baker, Can. Ent., xxxix, p. 115, 1907. 

The original description follows: 

Similar in size and form to coquilletti Coding; the markings are less promi- 
nent, lateral horns much less produced. 

Male. Greenish yellow, marked with ferruginous lines, punctured. Head 
greenish j'ellow, lightly punctured. Prothorax yellowish green, mottled with 
ferruginous, longitudinal, elevated lines; dorsal carina percurrent, a deeply 
impressed dot on each side of its base; dorsal crest somewhat elevated, much 
compressed, strongly compressed anteriorly at base and posteriorly behind 
middle, the highest point of crest at beginning of postesior third, from which 
point it gradually slopes anteriorly in a gentle curve continuous with anterior 
third of prothorax, posteriorly sloping for a short distance, then forming an 
obtuse angle; at the base another obtuse angle is formed, from which the 
median carina curves gently to the apex; lateral angles a little prominent; 
tegmina with basal half coriaceous, apical half subcoriaceous, a brown spot at 
apex. Chest below is dark yellow, coxae piceous. Legs yellow and hairy; 
abdomen yellow. Length, 6 mm. 

Habitat: Mario county, Cal. 



78 The University Science Bulletin. 

Distribution. The type specimen came from California. There 
are specimens in the Snow collection from Arizona and from Lincoln 
county, Kansas. 

Hosts. Unknown. 

Telamonanthe modesta (Goding). 

Telamoim modesta Goding, Bui. III. St. Lab. Nat. Hist., iii, p. 420, 1894. 

The original description follows: 

Head triangular, hairy; ocelli nearer to each other than to the eyes. 
Prothorax broad, convex in front, gradually elevated back of lateral angles in 
a very high, much compressed, crest, the upper and anterior edges continuously 
curved to base of prothorax; posterior superior angle rectangular, posterior 
edge straight, inclined forward somewhat; posterior process long, depressed, 
acuminate, gradually attenuated to apex; sordid greenish yellow covered with 
black punctures, hairy, two black impressed dots over each eye, one above the 
other; base of posterior process and posterior edge of crest more or less free 
from black punctures. Tegmina with basal half of corium punctured, sub- 
transparent. Legs triquetrous, tibiae punctured with black, covered with 
spines. Abdomen and chest greenish-yellow. 

Length. 8 mm.; breadth 4 mm.; altitude, 5 mm. 

Habitat : Galesburg, 111. 

Distribution. Types from Illinois. There is a specimen in the 
Snow collection from Kansas City, Mo., so the species undoubtedly 
occurs in Kansas. 

Hosts. Unknown. 

Genus Archasia Stal. 

The members of this genus are at once recognized by the broad, 
leaflike and compressed expansion of the pronotum. 

Van Duzee separates the two species occurring in Kansas by the 
following key: 

* KEY TO SPECIES. 

A. Dorsal edge distinctly brown or fuscous, the contour obviously concave 
before the apex. belfragei. 

AA. Dorsal edge concolorous or faintly dotted with brown, contour scarcely 
if at all concave before the apex. galeata. 

Archasia belfragei Stal. 

Archasia belfragei Stal, Of. Vet. Akad. Forh., xxvi, p. 250, 1869. 
Archasia canadensis Piovancher, Pet. Faune Ent. Can., iii, p. 230, 1889. 

Funkhouser gives the following technical description: 

Green, fading to yellowish in cabinet specimens; pronotum high, strongly 
foliaceous, dorsal margin brown; tegmina about half concealed by pronotum; 
posterior process not reaching apices of tegmina. 

Head nearly twice as wide as long, smooth, sparingly' pubescent; base high 
and sinuate; eyes very prominent, shining dark brown; ocelh pearly, promi- 
nent, nearer to each other than to the eyes. 



Lawson: Membracid.e of Kansas. 79 

Pronotuni cioscly bvit weakly piinctatp, not pubescent; humeral angles 
small, triangular; dorsal crest very high, flattened, foliaceous, almost vertical 
above head, slightly concave above head, posterior margin gradually hollowed 
out before apex of posterior process, entire dorsal margin flattened and uni- 
formly brown. 

Tegmina smoky iiyaline, bases and costal margins punctate, tips strongly 
marked with brown. Understirface of body yellow-brown; abdomen brown 
Legs dull yellow-brown; tibiae pubescent. 

Length, 9 mm.; width, 4.5 mm.; height of pronotum, 5 mm. 

Distribution. Van Duzee reports this species from Ontario, 
Massachusetts, New York, New Jersey, North Carolina, Illinois and 
Michigan. There is a specimen in the Snow collection from Kansas 
City, Mo., so it undoubtedly occurs in Kansas also. 

Hosts. Funkhouser gives locust and oak as hosts. 

Archasia galeata (Fabricius.) 

Mefnbracis galeata Fabricius, Syst. Rhyng., p. 9, 1803. 

Thelia yaleata Fairniaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 309, 1846. 

Smilia auriciUata Emmons, Nat. Hist. N. Y. Ins., p. 153, pi. 3, fig. 12, 1854. 

The following describes this species: 

Green, fading to yellowish in cabinet specimens; pronotum very high and 
strongh' foliaceous, dorsal margin concolorous or spotted with brown; tegmina 
about half concealed by pronotimi ; posterior process not reaching apices of 
tegmina. 

Head shghtly longer than in preceding species, smooth, sparingly pubescent; 
base high and sinuate; eyes very prominent, shining dark brown; ocelli pearly, 
prominent, nearer to each other than to the eyes. 

Pronotum closely but distinctly punctate, sparsely pubescent; humeral an- 
gles smaller and more rounded than in preceding species and the dorsal crest 
higher, not concave above the head, with posterior margin not at all concava 
before apex of posterior process, the entire dorsal margin strongly flattened. 

Tegmina smoky hyaline, bases and costal margins punctate, tips slightly 
darker. Entire undereurface and legs yellowish-brown, the tibiae pubescent. 

Length, 9-11 mm.; width, 4.5-5 mm.; height of pronotum, 6 mm. 

Distribution. Van Duzee reports this species from Ontario, New 
York, New Jersey, Pennsylvania, North Carolina, Georgia, Florida, 
Illinois, Iowa, Colorado, Utah and Texas. Specimens are at hand 
from Douglas, Riley, Miami and Cowley counties, Kansas. 

Hosts. Coding mentions Eupatorium, Verbena hastata and oak. 

Genus Smilia Germar. 

The members of this genus are of the same general shape as those 
of the preceding, because of the foliaceous dorsum, but the terminal 
cell of the hind wing is not sessile and truncate, but petiolate and 
triangular. 

A single species occurs in Kansas. 



80 The University Science Bulletin. 

Smilia camelus (Fabricius). 

(PI. Vll, figs. 1, 2.) 

Membracis camelus Fabricius, Syst. Rhyng., p. 10, 1803. 

Smilia vittata Amyot & Sendlle, Heniip., p. 539, 1843. 

Thelia camelus Walker, List Homop., ii, p. 562, 1851. 

Thelia vittata Walker, List Homop., iv, p. 1143, 1852. 

Smilia betulae Coding, Can. Ent., xxv, p. 196, 1893. 

Antianthe compressa Buckton, Monog. Memb., p. 191, pi. 41, fig. 6, 1903. 

. Funkhouser gives the following technical description : 

Pronotimi high and foliace'ous, extending forward over the head; brown 
with broad diagonal stripe of green or. yellowish followed by a parallel trans- 
lucent band and a white spot; males much smaller and darker than females. 

Head triangular, sculptured, yellow with scattered brown punctures and 
hairs; eyes brown; ocelli pearly, margins raised, nearer to each other than 
to the eyes; clypeus continuing inferior line of face, apex slightly produced. 

Pronotum coarsely punctiu-ed, punctures farther apart in pale parts; wide 
green band extending from anterior dorsal angle of crest to lateral margin 
of pronotum, this band fading to yellowish in dried insects; wide translucent 
band from behind middle of dorsum to lateral base of crest; white spot at 
posterior base of crest; humeral angles hardly produced, short, rounded; 
posterior process short, pointed, not reaching tips of tegmina. 

Tegmina hyaline, bases punctate with brown, apices brown. Undersur- 
face of body brownish yellow. Legs flavous. 

Length: female, 9 mm.; male, 7-8 mm. Width: female, 3 mm.; male, 
2.5-3 mm. 

Internal male genitalia. Styles with anterior portions broadened 
and flattened as in Telamona, the sparsely spined posterior portions 
of nearly same width to the laterally curved and doubly toothed 
apices; connective large, broad, much as in Telamona; oedagus, 
viewed laterally, large, U-shaped, anterior arm lightly chitinized 
and longer than the more heavily chitinized posterior arm, with a 
distinct knob at base and also about midway up the anterior arm, 
the posterior arm with functional orifice on caudal aspect of apex 
and covered with teeth on the cephalic aspect. 

Distribution. Van Duzee reports this species from Ontario, New 
Hampshire, Massachusetts, New York, New Jersey, Pennsylvania, 
North Carolina, Georgia, Florida, Illinois, Michigan, Iowa, Mis- 
souri, and Texas. Specimens have been taken in Kansas in Douglas 
and Montgomery counties. 

Hosts. Funkhouser gives locust and oak as hosts. 

Genus Cyrtglgbus Goding. 
The members of this genus are usually brownish insects without 
prominent humeral angles, with a compressed dorsum, and fre- 
quently with a thin, semitransparent spot below the dorsal ridge. 



Lawson: Membracid.e of Kansas. 81 

Tlie genus is represented by at least nine species in Kansas, which 
live on various species of oaks. 

Van Duzee divides the genus into four subgenera which he sepa- 
rates by the following key : 

KEY TO SUBGENERA. 

Pronotum strongly inflated posteriorly, the crest forming an inflated cyst be- 
fore and behind the median pale spot. Xantholobus. 

Pronotum not strongly inflated, the crest however, usually more or less dis- 
tinctly compressed on the median spot 1 

1. Dorsal crest low and sinuated at the middle; form elongated and 
much depressed. Evashmeadea. 
Dorsal crest not distinctly sinuated above 2 

2. Crest highest anteriorly over or before the humeral angles, the dor- 
sum sloping and nearly rectilinear to the apex. Atymna. 
Crest regularly arcuated above in the more typical examples, or in 
any case with the highest point at or near the middle. 

Cyrtolobus. 

Subgenus Cyrtolobus Coding. 

KEY TO SPECIES. 

A. Dorsum without anterior notch or depression. 

B. Crest well developed. celsiis. 

BB. Crest distinctly lower. fenestratus. 

AA. Dorsum with anterior depression before elevation. 

B. Crest arising before humeral angles. fuliginosus. 

BB. Crest arising behind humeral angles. 

C. Large species, at least 9 mm. in length. tuberosus. 

CC. Small species, not over 7 mm. in length. 

D. Crest low or obsolete. 

E. Species distinctly brownish. griseus. 

EE. Species distinctly greenish. cinereus. 

DD. Crest well developed; pronotum with distinct oblique 
bands. vau. 

Cyrtolobus celsus Van Duzee. 

Cyrtolobus celsus Van Duzee, Check List Hemip., p. 60, 1916 (n. n. for fenestratus 
Van D.). 

Cyrtolobus fenestratus Van Duzee, Bui. Buf. Soc. Nat. Sci., ix, p. 81, 1908. 

The following is the original description: 

Pronotum well elevated, highest at about the middle, almost regularly arcu- 
ated from base to tip, the dorsal line a very little broken at the posterior vitta. 
Head and the pronotum anterior to the oblique line pale yellowish testaceous, 
punctured and varied with brown; from above each eye a brown indefinite 
vitta curv^es backward over the humeral angle; anterior oblique vitta almost 
perpendicular at first, approaching the elongated median mark, then deflected 
and again widened at apex so as to pass almost straight across the dorsal 
carina ; posterior vitta transverse, represented in my Atlanta specimen only by 
a narrow spot on the carina; the surface behind the anterior vitta darker, in 
one example almost piceous. Elytra hyaline, the smoky apex small and palo. 



82 The University Science Bulletin. 

Face coarsely punctured; clypeus broad, moderately produced, and strongly 
incurved. 

Length, 6 mm.; height, 3 mm. 

Distribution. Van Duzee reports this species from Massachu- 
setts, New York, New Jersey and Georgia. It has also been taken 
near Kansas City, Mo., and therefore undoubtedly occurs in Kansas. 

Cyrtolobus fenestratus (Fitch). 

Cyrtosia fenestrata Fitch, Hoinop. N. Y. St. Cab., p. 49, 1851. 

Cyrotolobus fenestratus Goding, Can. Ent., .xxv, p. 172, 1893. 

Cyrtolobus muticus Van Duzee, Bui. Buf. Soc. Nat. Sci., ix, p. 83, 1908 (in part). 

The following is the original description: 

Yellow marbled with rufous; a pellucid spot behind the summit of the keel 
and a smaller one halfway to the apex; an oblique yellow vitta below the an- 
terior spot, margins with fuscous or sanguineous; tip of the thorax reaching 
beyond the terminal cells of the elytra. Male black, the pellucid spots almost 
obsolete and the yellow vitta replaced by a few yellow dots. 

Length, 0.25. On oaks. 

Distribution. Van Duzee reports this species from Massachu- 
setts, New York, New Jersey, Pennsylvania, North Carolina, Geor- 
gia, Florida, Mississippi, Ohio, Illinois, Colorado, Dakota. Speci- 
mens have been taken at Kansas City, Mo., so it surely occurs in 
Kansas. 

Cyrtolobus fuliginosus (Emmons). 

Cyrtosia fuliginosa Emmons, Nat. Hist. N. Y. Ins., p. 154, pi. 13, fig. 15, 1854. 
Cyrtolobus fuliginosus Goding, Can. Ent., xxv, p. 172, 1893. 

Funkhouser gives the following technical description: 

Near C. ovatus in appearance, but smaller, darker, and with lower crest; 
dark sordid brown with faint transverse bands; head projecting slightly for- 
ward; posterior process just reaching tips of tegmina; tegmina strongly 
marked with brown, apices lighter. 

Head somewhat extended forward, yellow, mottled with deep brown, deeply 
punctate with brown, not pubescent, a black spot at base of head above each 
ocellus; eyes large, brown, lighter in color than remainder of head; ocelli 
small, pearly, about equidistant from each other and from the eyes; clypeus 
convex, sculptured, a brown line on each side, tip continuing rounded inferior 
outline of face. 

Pronotum dark brown, transverse fascia extending from anterior base of 
crest to lateral margin of pronotum, this fascia light browTi before and very 
dark brown behind; entire pronotum deeply and densely punctate; humeral 
angles weak, angular but blunt; dorsal crest regularly arcuate from above 
humeral angles to base of posterior process; posterior process heavy, short, 
blunt, just reaching apices of tegmina. 

Tegmina smoky brown, apical cells lighter, apical margins fuscous, bases 
and costal margins roughly punctate. Legs and undersurface of body flavous. 

Length, 6 mm.; width, 2.5 mm. 



Lawson: Mkmbracid.e of Kansas. 83 

Distribution. Hitherto reported only from New York. Specimens 
have been taken, however, at Kansas City, Mo., and so it may safely 
be included in the Kansas fauna. 

Hosts. Funkhouser reports it from white oak. 

Cyrtolobus tuberosus (Fairmaire). 

Thelia tuberosus Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 307, 1846. 
Cyrtolobus tuberosus Goding, Bui. III. St. Lab. Nat. Hist., iii, p. 433, 1894. 

Funkhouser gives the following technical description: 

Largest species of the genus; brown mottled with darker brown; dorsal 
compression strikingl}- transparent; dorsal crest situated well back on prono- 
tuni. posterior process very short; tegmina smoky hyaline tipped with brown. 

Head triangular, broader than long, ochraceous tinged with red and punc- 
tate with brown, not pubescent; base weakly sinuate; inferior margin of face 
strongly sinuate; eyes large, brown; ocelli small, yellowish, slightly protrud- 
ing, nearer to each other than to the eyes; clj'peus convex, brown line on each 
side, tip extended and hairy. 

Pronotum deeply and closely punctate, light greenish brown; crest dark 
brown with pale compression at anterior base, in the middle, and at posterior 
base; middle compression veiy large and transparent, posterior half of crest 
dark brown with color extending in a dark band to margin of pronotum; 
metopidium very convex, median carina prominent; humeral angles promi- 
nent, rounded; posterior process short, sharp, brown, inferior lateral margin 
slightly sinuate, not reaching tips of tegmina. 

Tegmina brownish h3-aline, tips strongly marked with brown, bases punc- 
tate. Undersurface of thorax yellow. Legs ferruginous, hind trochanters 
marked with brown; tarsi flavous; claws brown. 

Length, 9.5 mm.; width, 4 mm. Male smaller than female, but similarly 
colored. 

Distribution. Reported by Van Duzee from Massachusetts, New 
York, North Carolina, Georgia, Florida, Mississippi and Illinois. 
There is a specimen in the Snow collection from Kansas City, Mo., 
so it is sure to occur in Kansas. 

Hosts. Funkhouser reports it from red oak and hickory. 

Cyrtolobus griseus Van Duzee. 

Cyrtolobus griseus Van Duzee, Bui. Buf. Soc. Nat. Sci., ix, p. 90, 1908. 

The following is the original description : 

This form, which is very near ciner'eus, is of an almost uniform dark cinerous, 
closely punctured with fuscous. The only conspicuous marking is the oblique 
blackish vitta which in cinercus bounds the hind margin of the anterior oblique 
vitta. Before and behind this brown band or line the surface is a little lighter 
and there is a suggestion of the median dorsal spot and transverse posterior 
vitta. In some specimens there is an arcuated longitudinal brown vitta on 
either side of the metopidium. Here the elytra are hyaline with the costal 
base ferruginous grey and punctured, and the apex has a small fuscous cloud. 



84 The University Science Bulletin. 

The tergum is black with a broad white band at the base as in the alHed spe- 
cies, which, showing through the closed elytra, indicates the markings found 
on the elytra of cinereus. Face obviously longer and more convex than in 
cinereus, with the basal middle depressed, and the clypeus and lorse together 
larger, the former much broader, more convex and more decurved and rounded 
at apex. 

Distribution. The type material was taken at Effingham, Kansas. 

Cyrtolobus cinereus (Emmons). 

Gargara cinereus Emmons, Nat. Hist. N. Y. Ins., p. 156, pi. 13, fig. 3, 1854. 

Cyrtosia ornata Provancher, Pet. Faune Ent. Can., iii, p. 240, 1889. 

Cyrtosia cinerea Harrington, Ottawa Nat., vi, p. 30, 1892. 

Cyrtolobus cinereum Coding, Can. Ent., xxv, p. 172, 1893. 

Atymna cinereum Coding, Bui. 111. St. Lab. Nat. Hist., iii, p. 436, 1894. 

Cyrtolobus cinereus Van Duzee, Bui. Buf. Soc. Nat. Sci., ix, p. 91, 1908. 

Funkhouser gives the following technical description: 

Small greenish gray mottled with brown and banded with green; pronotum 
low and regularly arcuate; metopidium convex; posterior process short but 
sharp; tegmina wrinkled, hyaline, apices brown. 

Head convex, pale grayish green, sharply punctate with black, sparingly 
pubescent; base nearly straight; eyes prominent, brown; ocelU large, reddish, 
prominent, slightly farther from each other than from the eyes and situated 
sUghtly below an imaginary line extending through centers of eyes; clypeus 
flat, somewhat trilobed, a faint brown line on each isde, extending below in- 
ferior margin of face. 

Pronotum green-gray tinged with reddish, closely punctate, not pubescent; 
dorsal crest very low, median spot on margin pale ; a transverse pale band bor- 
dered with brown extending from anterior base of crest backward and down- 
ward to lateral margin of pronotum, a similar band extending from base of 
posterior process downward and forward to almost meet the anterior stripe 
and form a V with it ; posterior process short, not reaching tips of tegmina. 

Tegmina wrinkled, hyaline, brown spot at base of each, another in middle, 
and a third at tip; areas between hyaline. Legs and undersurface of body 
grayish flavous. 

Length, 5.8 mm.; width 2.5 mm. 

Distribution. Reported by Van Duzee from Quebec, New York 
and New Jersey. A single specimen of this species was taken in 
Douglas county, Kansas, and is in the Snow collection. Specimens 
have also been taken in Riley county. 

Cyrtolobus i^au (Say). 

(PI. Vll, figs. 5, 6.) 

Membracis van Say, Jl. Acad. Nat. Sci. Phila., vi, p. 299, 1831. 

Thelia seinifascia Walker, List Homop., ii, p. 561, 1851. 

Smilia vau Fitch, Homop. N. Y. St. Cab., p. 48, 1851. 

Thelia vau Walker, List Homop., iv, p. 1142, 1852. 

Cyrtosia vau Provancher, Pet. Faune. Ent. Can., iii, p. 238, 1889. 

Cyrtosia jenestrata Provancher, Pet. Favme Ent. Can., iii, p. 239, 1889. 

Cyrtolobus nigra Coding, Can. Ent., xxv, p. 172, 1893. 

Cyrtolobtis punctifrontis Coding, Can. Ent., xxv, p. 172, 1893- 



Lawson: Membracid^ of Kansas. 85 

Cyrtolobus tricincta Coding, Can. Ent., xxv, p. 172, 1893. 

Cyrtolobus van Coding, Can. Ent., xxv, p. 172, 1893. 

Thelia fasciata Buckton, Monog. Membr., p. 189, 1893. 

Argante semifasciata Buckton, Monog. Membr., p. 189, pi. 40, fig. 9, pi. 41, fig. 1, 1903. 

Cyrtolobus variM Smith, Cat. Ins. N. J., edn. 3, p. 92, 1910. 

Funkhouser gives the following technical description: 

Small robust species, with low pronotum and prominent markings; varies 
greatlj' in color and somewhat in size; females larger and lighter than males, 
but with constant markings; transverse pronotal band prominent, pale bor- 
dered with deep brown; dorsal compression deep and translucent; posterior 
process short, blunt, not reaching tips of tegmina; tegmina hyaline, with bases 
and tips slightly brown. 

Head small, subtriangular, pale yellow punctured with brown; base feebly 
sinuate; inferior margin of face sinuate, c^'peus extending slightly below line; 
eyes large, gray-brown; ocelli small, yellowish, somewhat nearer to each other 
than to the eyes; clypeus hairy. 

Pronotum closely and roughly punctate; median compressed spot round, 
transparent; dorsal crest low, arising above humeral angles and gradually ex- 
tending with only a faint sinus before posterior process; posterior process short, 
blunt, tectiform, reaching to bases of apical cells of tegmina. 

Tegmina hyaline, veins prominent, bases and apices smoky hyaline. Legs 
and undersurface of body uniform flavous. 

Length, 5.5-6.5 mm.; width, 2.4-2.6 mm. 

Internal male genitalia. Styles rather long and slender, curved 
strongly laterad apically, and bearing several spiny tubercles on the 
lateral margins just cephalad of the terminal tooth; connective tri- 
angular; cedagus, viewed laterally, large, U-shaped, anterior arm 
slightly chitinized and with a large prominence about half way on 
its cephalic aspect, posterior arm heavily chitinized, apex with file- 
like teeth on the side opposite the functional orifice. 

Distribution. Reported by Van Duzee from Ontario, Maine, New 
Hampshire, Massachusetts, New York, New Jersey, Pennsylvania, 
District of Columbia, North Carolina, Georgia, Florida, Illinois, 
Kansas, Missouri, Arkansas, Texas, New Mexico and Colorado. 
There are specimens in the Snow collection from Douglas, Riley, and 
Pottawatomie counties, Kansas. 

Hosts. Funkhouser gives white, chestnut, red and scarlet oak 
as hosts. 

Subgenus Atymna Stal. 

In this subgenus the pronotum is highest at the anterior end. 
It is represented in Kansas by a single species. 

Cyrtolobus querci (Fitch). 

(PI. VII, figs. 3, 4.) 

Smilia querci Fitch, Homop. N. Y. St. Cab., p. 49, 1851. 

Thelia querci Walker, List Homop., iv, p. 1143, 1852. 

Gargara querci Emmons, Nat. Hist. N. Y. Ins., p. 156, pi. 13, fig. 8, 1854. 



86 The University Science Bulletin. 

Atymna querci Van Duzee, Psyche, v, p. 390, 1890. 

Cyrtolohus {Atymna) querci Van Duzee, Bui. Buf. Soc. Nat. Sci., v, p. 188, 1894. 

Funkhouser gives the following technical description: 

Females large and green, males smaller and brown with a broken yellow 
median dorsal stripe; body long and narrow; crest highest above humeral 
angles and gradually sloping to posterior apex without a sinus. 

Female. Head projecting slightly forward, pale yellow, sculptured, irregu- 
larly punctate, not pubescent; eyes very prominent, reddish; ocelli not promi- 
nent, yellow; clypeus extending below inferior margin of face. 

Pronotum uniform green, roughly punctate, not pubescent, dorsal line 
faintly marked with brown; posterior process short, acute, not reaching tips 
of tegmina. 

Tegmina entirely hyaline, bases and costal margins faintly punctate; hind 
wings iridescent. Legs and imdersurface of body green. 

Length, 7 mm.; width, 2.5 mm. 

Male. Head sordid yellow, sculptured, sparingly punctate; eyes prominent, 
brown; ocelli pearly; clypeus marked with brown at base. 

Pronotum chocolate brown with bright yellow stripe on median dorsal line 
and yellow band before apex. 

Tegmina smoky hyaline with brown cloud at apices. Undersurface of 
thorax brownish; abdomen xevy dark brown, nearly black. Legs flavous; tarsi 
ferruginous; claws fuscous. 

Length, 6 mm.; width, 2 mm. 

Internal male genitalia. Styles small, anterior portion quite short, 
posterior portions larger, bearing spiny protuberances apically ; con- 
nective rather large, triangular; oedagus, viewed laterally, very large 
compared with the styles, anterior arm long and slender, posterior 
arm stout and with many filelike teeth on the side opposite the 
functional orifice. 

Distribution. Van Duzee reports this species from Ontario, Con- 
necticut, New York, Pennsylvania, North Carolina, Georgia, Illi- 
nois, Michigan, Iowa and Missouri. Specimens having been taken 
at Kansas City, Mo., it is sure to occur in Kansas. 

Subgenus Xantholobus Van Duzee. 

In this subgenus the pronotum is distinctly inflated posteriorly. 
It is represented in Kanas by a single species. 

Cyrtolohus muticus (Fabricius). 

Membracis mutica Fabricius, Genera Ins., p. 297, 1776. 

Cicada mutica Gmelin in Linnaeus, Syst. Nat., edn. 13, i, pt. 4, p. 2093, 1778. 

Centrotus mutica Fabricius, Syst. Rhyng., p. 21, 1803. 

Membracis trilineatus Say, Long's 2nd. Exped., p. 300, 1824; Compl. Writ., i, p. 200. 

Cyrtosia mutica Stal, Heniip. Fabr., ii, p. 25, 1869. 

Cyrtosia trilineata Provancher, Pet. Faune Ent. Can., iii, p. 239, 1889. 

Cyrtolobus muticus Coding, Bui. 111. St. Lab. Nat. Hist., iii, p. 431, 1894. 



Lawson: Membracid^ of Kansas. 87 

Funkhouscr gives the following technical description: 

Yellowish tinged with red; transverse band of pronotuni often absent; 
pronotum long; head slightly projecting forward; eyes tinged with reddish; 
posterior process reaching tijxs of tegniina ; teginina entirely hyaline or 
faintlj' clouded with yellow. 

Head slightly protruding forward, j-ellow with red punctures, sculptured, 
not pubescent; base irregularly sinuate; eyes gray marked with red; ocelli 
small, translucent, somewhat nearer to each other than to the eyes; clypeus 
swollen, convex, continuing inferior outline of face, tip slightly extended, 
hairy; antenme prominent. 

Pronotum yellowish with irregular reddish areas, deeply and roughly 
punctate, not pubescent; transverse band when present pale with reddish 
borders; humeral angles weak, blunt; dorsal crest elliptical, very slight sinus 
before base of posterior process, compressions not deep; posterior process 
heavj-, blunt, just reaching tips of tegmina. 

Tegmina hyaline or clouded with reddish yellow, tips pale, veins in some 
cases yellowish, bases and costal areas irregularly punctate. Legs and under- 
surface of thorax flavous; abdomen sordid yellow. 

Length, 6 mm.; width, 2.8 mm. 

Distribution. Reported hy Van Duzee from Quebec, Rhode 
Island, New York, Pennsylvania and Illinois. In the Snow col- 
lection there are specimens from Kansas City, Mo., and some taken 
in Douglas county, Kansas. The writer took a specimen at St. 
Paul, ]\Iinn. 

Genus Ophiderma Fairmaire. 

Tlie members of this genus have a compressed and rounded dor- 
sum which entirely lacks a crest. 

It is represented in Kansas by three species, which may be sep- 
arated by the following key: 

KEY TO SPECIES. 

A. Color, browTi or mottled. 

B. Species smaller, mottlings more distinct. salamandra. 

BB. Species larger, mottling dull. flaviguttula. 

AA. Color, green or yellowish green. flava. 

Ophiderma salamandra Fairmaire. 

(PI. VI, figs. \ 2.) 
Ophiderma salamandra Fairmaire, Ann. Soc. Ent. Fr., ser. 2, iv, p. 493, 1846. 

Funkhouser gives the following technical description: 
Large brown species; dorsum rounded and very pubescent with short, black, 
bristly hairs; posterior process short, suddenly acute, not reaching apices of 
tegmina; tegmina hyaline, bases and costal areas strongly punctate, tips 
clouded with fuscous, veins very prominent ; under part of body dark ; males 
smaller and darker than females. 



88 The University Science Bulletin. 

Head broader than long, yellow, feebly punctate, very hairy; base slightly, 
uniformly curved; eyes large, bro^vn; ocelli prominent, red, nearer to each 
other than to the eyes; inferior margin of face sinuate; clypeus yellow with 
two vertical stripes of red; base hairy. 

Pronotum coarsely punctate, densely pubescent, brown mottled with green; 
dorsum rounded, slightly depressed behind middle, lateral margin curved 
downward at middle; posterior process short, suddenly acute, not reaching 
tips of tegmina. 

Tegmina smoky hyline, veins very prominent, nearly all of basal half below 
pionotum strongly punctate, tips clouded with fuscous; hind wings iridescent. 
Undersurface of head and thorax fuscous; abdomen flavous. Femora and 
tibiae strongly marked with dark brown. 

Length, 7.6 mm.; width, 3.2 mm. 

Internal male genitalia. Styles rather short and broad, especially 
basally, posterior portion more slender, apices curved strongly 
laterad and ending in a small but distinct terminal hook on the 
lateral margin, in front of which is a distinct prominence which 
tapers gradually to the hook, the apical fourth of the styles bearing 
a few spines; connective almost pentagonal, with a pair of distinct 
knobs at the basal angles; oedagus, viewed laterally, U-shaped, 
anterior process with a large anterior prominence near the middle, 
posterior arm curved, the apex bearing the functional orifice caudally 
and many filelike and distinct teeth on its cephalic aspect. 

Distribution. Reported by Van Duzee from Ontario, New Hamp- 
shire, Massachusetts, New York, New Jersey, Pennsylvania, District 
of Columbia, Virginia, Georgia, Florida, Illinois and Michigan. 
There are specimens in the Snow collection also from Wisconsin, 
Arizona and Kansas City, Mo. It has also been taken in Potta- 
watomie county, Kansas. 

Hosts. Oaks. 

Ophiderma jiaviguttula Coding. 

Ophiderma jiaviguttula Coding, Bui. 111. St. Lab. Nat. Hist., iii, p. 438, 1894. 
Ophiderma flavoguttata Slosson, Ent. News, xvii, p. 326, 1906. 

The original description follows: 

Female. Head triangular, yellowish; eyes prominent, dark brown; ocelli 
equidistant from each other and from the eyes, red; convex, densely pubes- 
cent. Prothorax with very slight median carina, densely pubescent, an 
irregular yellow patch starting at lateral border and extending upwards and 
forwards, midway between base and apex; an irregular band at base, con- 
colorous with head, extending along sides in a greenish gray line; otherwise 
dirty brown, lightly punctvired; apex of posterior process not reaching apex 
of tegmina. Tegmina' subcoriaceous at base, lightly punctured, basal half 
and apex brown. Below yellow, feet and legs brown. 

Length, 6.2 mm. 



Lawson: Membracid^ of Kansas. 89 

Distribution. Reported by Van Duzee from New Hampshire, 
Pennsylvania and Illinois. There arc specimens in the Snow col- 
lection from Kansas City, Mo., so it undoubtedly occurs in Kansas. 

Hosts. Probably oak. 

Ophiderma fiava Goding. 

Ophiderma fiava Goding, Insect Life, v, p. 93, 1892 (nomen nudum). 
Ophiderma fiava Goding, Bui. 111. St. Lab. Nat. Hist., iii, p. 439, 1894. 

Funkhouser gives the following technical description: 

Large greenish yellow species, fading to sordid yellow in cabinet specimens; 
body robust and long; posterior process not reaching apices of tegmina; 
tegmina h3'aline, brown at base and fuscous-clouded at tips. 

Head much broader than long, green, weakly and sparingly punctate, 
smooth, shining, sparingly pubescent; eyes large, red; ocelli prominent, red- 
dish, about equidistant from each other and from the eyes; clypeus smooth, 
nearh^ black, base regularh' rounded, tip extending below inferior margin of 
face. 

Pronotum uniform green, in some cases tinged with reddish, closely and 
densel}^ punctate, finely pubescent; dorsum rounded, depressed behind middle, 
median carina percurrent; posterior process heavy, tectiform, acute, not ex- 
tending to tips of tegmina. 

Tegmina hyaline, bases reddish and punctate, tips clouded with fuscous, 
veins heavy and inclined to be punctate along margin. Legs and undersur- 
face of body entirely flavous. 

Length, 7-8 mm.; width, 3-4 mm. 

* Distribution. Reported by Van Duzee from Quebec, Massa- 
chusetts, New York, Pennsylvania, North Carolina, Illinois, Michi- 
gan. There are specimens in the Snow collection from Columbia, 
Mo., and from Chautauqua county, Kansas. 
Hosts. Unknown. 

Genus Vanduzea Goding. 
The members of this genus are distinguished by the transverse 
and basally truncate terminal cell of the elytra. 
A single member of the genus is known to occur in Kansas. 

Vanduzea triguttata (Burmeister). 

(PI. VI, figs. 7, 8.) 

Entylia triguttata Burmeister, Silb. Rev. Ent., iv, p, 183, 1836. 

Vanduzea vestita Goding, Insect Life, v, p. 93, 1892. 

Cyrtolobus annexus (Uhl. MS) Townsend, Can. Ent., xxiv, p. 196, 1892. 

The following is Coding's description : 

Head broad, black, perpendicular, triangular, a narrow dusky brown mark 
on upper edge just below origin of carina. Eyes prominent; ocelli equidistant 
from each other and the eyes. Front of prothorax blackish brown, fading 
posteriorly to a reddish brown in a triangular form, the apex of which reaches 



90 The University Science Bulletin. 

three-fourths of the distance to apex of posterior process ; lateral angles slightly 
produced; sides of prothorax from just behind lateral angles to apex black, 
interrupted by a light yellow, or whitish, trapezoidal spot on each side just 
behind middle of inferior border; just before the apex a white band across 
posterior part of process. Tegmina clear, with dark brown veins, or brown 
with a lighter band across middle. Legs and feet brown or .black. 
Length, 4.7 mm. 

Internal male genitalia. Styles broad and flat basally, apical half 
slender, apices curved strongly dorsad and ending rather bluntly, 
bearing a few spines ; connective nearly quadrangular, the base con- 
cave; oedagus, viewed laterally, U-shaped, anterior arm enlarged 
toward the base, posterior arm of nearly same width throughout, 
the apex bearing caudally the functional orifice and cephalad many 
filelike teeth. 

Distribution. Reported by Van Duzee from District of Columbia, 
Florida, Colorado, New Mexico and Arizona. The writer has taken 
it at St. Paul, Minn. It has been taken in the following Kansas 
counties: Morton, Clark, Stevens, Logan, Haskell, Seward, Riley, 
Hamilton, Pottawatomie, Douglas and Miami. 

Hosts. Common on black locust. Also taken by the writer on 
A7norpha. 

Genus Entylia Germar. 

The members of this genus are characterized by their high com- 
pressed dorsum, which bears a deep median notch. 

A single species of the genus is known to occur in Kansas. 

Entylia concisa Walker. 

(PI. VI, figs. 5, 6.) 

Entylia concisa Walker, List Homop., ii, p. 547, 1851. 

Entylia decisa Walker, List Homop., ii, p. 548, 1851. 

Entylia concava Piovancher, Pet. Faune Ent. Can., iii, p. 233, 1889. 

The following is the original description: 

Ferruginea; prothoracis carina alte bicristata, utrinque albo interrupte et 
oblique fasciata ; pedes flavi ; alse limpidfe ; alse anticse basi et ad costam fulvae. 

Ferruginous; head and thorax roughly punctured; head transverse, almost 
semicircular, narrower than the fore-chest, slightly impressed with an indis- 
tinct middle suture which extends to the face, the hind border of the latter is 
angular and occupies nearly half the length of the head; shoulders very ob- 
tusely angular, not prominent: fore-chest forming two lofty compressed keel- 
shaped crests which inchne towards each other and inclose three-fourths of a 
circle; the first rises between the shoulders and is truncated at the tip; the 
second Is lower and above the keel; the latter is rather deep and extends far 
beyond the tip of the abdomen, whose sides it embraces; the irregular ridges 
on the sides of the crest communicate with the ridges of the keel, and the lat- 
ter has an obUque white interrupted band on each side behind the second 



Lawson: Membracid.e of Kansas. 91 

crest; breast jiitchy; abdomen black; a stripe on each side beneath and the 
tip yellow ; legs yellow ; wings colorless ; fore-wings tawny at the base and along 
more than half the length of the fore border; veins pitchy, tawny where the 
wings are so, pale yellow near the tip of the fore border. 

Length of the abdomen, 2% hnes; of the wings, 4% lines. 

Var. /)' . Breast and abdomen tawny; white bands of the keel hardly visible. 

Var. y. Breast and abdomen black; tip of the latter tawn3\ 

St. John's Bluff, E. Florida. 

Internal male genitalia. Styles small, anterior portion narrow, 
posterior part stouter, the apices curved strongly laterad and ending 
in a stout hook; connective large, triangular, apex abtuse, and longi- 
tudinally divided; oedagus, viewed laterally, U-shaped, anterior arm 
with a distinct protuberance, posterior arm stouter and ending in a 
large, serrated point. 

Distribution. Van Duzee reports this species from District of 
Columbia, North Carolina, Georgia and Florida. It has been taken 
in Douglas, Pottawatomie and Wilson counties, Kansas. 

Remarks. Funkhouser feels that E. sinuata is the very small 
southern form, E. bactriana the northeastern, and that E. concisa is 
our Kansas species. In the present confused status of the members 
of this genus it is perhaps best to accept his conclusions and to 
change our Kansas species to E. concisa instead of calling it E. sin- 
uata, as did Miss Branch. 

Hosts. Miss Branch reports this species from Melilotus alba, 
Cnicus altissinius, Phleum alpinum, Helianthus annuus, Medicago 
sativa, and Ambrosia sp. The writer has taken it very commonly in 
all its stages from Ambrosia triflda, Xanthium sp., and Helianthus 
tuberosus. 

Genus Publilia Stal. 

The members of this genus are closely related to those of the pre- 
ceding genus, but have a much lower crest and a much weaker me- 
dian notch. 

All of the members of the genus found in the United States occur 
in Kansas. These may be separated by the following key given by 
Van Duzee. 

KEY TO SPECIES. 

A. Dorsum straight or feebly arcuated, scarcely if at all sinuated; form more 
slender. mudesta. 

AA. Dorsum more elevated, obviously sinuated. 

B. Sides of the pronotum with longitudinal rugae which become more 
or less reticulated along the dorsum. concava. 

BB. Rugae of the pronotum strong, irregularly reticulated over its whole 
surface. reticulata. 



92 The University Science Bulletin. "^ ^^ 

Publilia modesta Uhler. 

(PI. VI, fig. 3, 4.) 

Publilia modesta Uhler, Bui. U. S. Geol. Geog. Surv., i, p. .344, 1876. 
Publilia bicinctura Coding, Ent. News, iii, p. 200, 1892. 

The original description follows: 

General form of P. concaua Say, but more decidedly vertical in front, and 
with the dorsal outline scarcely depressed before the middle; apex of the pro- 
notum more slender and acute. Color pale yellow; the head and fore part of 
pronotum clouded with pale brown; a spot above the humeri, a broad obhque 
band behind the middle, and a broad cloud on the apex grayish brown; the 
dorsal edge irregularly spotted with dark brown, and the oblique band sur- 
mounted by a large brown spot. The surface closely beset with series of coarse 
sunken jjunctures; the longitudinal and reticulated surface lines obsolete. 
Humeral margin of the sinus waved. Under side piceo-testaceous; the front, 
clypeus, pectu.s and venter, excepting the edges of the segments, black-piceous. 
Legs dull yellow, closely pubescent, clouded, and spotted with brown. 

Length, 4 to 4% mm.; breadth of pronotum, 2 mm. 

Colorado (C. Thomas) ; also discovered in Utah, Dakota, Arizona, New 
Me.xico, Cahfornia. , 

In two specimens examined, the commonly raised lines on the surface of the 
pronotum were obliterated. 

Distribution. Besides the above localities specimens have been 
taken in Kansas in Gove and Trego counties. 

Hosts. Gillette and Baker report this species on Solidago, alfalfa, 
Helianthus, Iva, and Artemesia. Goding gives Glycyrrhiza lepidota 
and mesquite as hosts. 

Publilia concava (Say). 

Membracis concava Say, Long's 2nd Exped., ii, p. 301, 1824; Compl. Writ., i, p. 200. 

Entylia concava Germar, Silb. Rev. Ent., iii, p. 249, 1835. 

PublUia concava Stal, .\nalecta Hem., p. 388, 1866. 

Ceresa concava Rathvon, in Mombert's Hist. Lane. Co., Pa., p. 5.51, 1869. 

Publilia grisea Buckton, Monog. Membr., p. 184, pi. 39, fig. 5, 1903. 

Publilia vittata Buckton, Monog. Membr., p. 185, pi. 39, fig. 6, 1903. 

Funkhouser gives the following technical description: 

Varies greatly in color and somewhat in shape, particularly in form of dor- 
sal sinuation; color varies from gray to black; dorsum convex, tectiform, 
faintly ribbed, dorsal sinus shallow; pronotum irregularly ridged, deeply punc- 
tate; tegmina largely co\ered by pronotum, basal half of each costal area 
strongly punctate. 

Head slightly projecting, strongly punctate with black; base nearly straight; 
inferior margin I'ounded; ej'es not prominent; ocelli prominent, usually red- 
dish; clypeus rounded, very wide at tip. 

Pronotum deeply, densely and coarsely punctate, lateral areas marked with 
high, distinct, irregular, longitudinal ridges; dorsal margin sinuate just be- 
hind humeral angles, sinuation usually very shallow; posterior lobe gradually 
elliptical to posterior apex; posterior process heavy, high, tectiform, blunt, ex- 
tending just beyond. tips of tegmina. 



Lawson: Membracid.e of Kansas. 93 

Tegmina almost entirely concealed by pronotum ; exposed costal margins 
opaque and punctate for basal half, ajncal areas hyaline, tips fuscous. Under- 
surface of body and femora usually very dark, generally black. Legs flavous. 

Length, 5 mm.; width, 2.5 mm. 

Internal male genitalia. Styles long and slender, the extreme an- 
terior portion bent strongly laterad, widest opposite connective, the 
posterior part with distinct apical hook; connective widest at the 
concave base, the apex truncate; oedagus, viewed laterally, with 
strong anterior arm which bears two prominent rounded promi- 
nences, the posterior arm more slender, the apical portion bearing a 
few indefinite teeth on caudal margin and the apex characteristically 
retrorsely hooked. 

Distribution. Van Duzee reports this species from Ontario, New 
York, New Jersey, North Carolina, Ohio, Illinois, Iowa, Missouri, 
Kansas, Arkansas, and Utah. It is also known to occur in Mexico. 
Specimens have been taken in Kansas in Douglas, Pottawatomie, 
Riley, and Gove counties. 

Hosts. Funkhouser gives golden rod and skunk cabbage as hosts. 
Goding mentions Canadian thistle. The writer has taken it in all 
stages on Ambrosia trifida. 

Publilia reticulata Van Duzee. 

Pxiblilia reticulata Van Duzee, Bui. Buf. Soc. Nat. Sci., ix, p. 106, 1908. 

The following is the original description : 

Closely allied to concava but with the surface of the pronotum reticulated 
with strong anastomosing rugae in place of the four or five simple longitudinal 
carinse found in that species. These rugae give the surface a strongly corru- 
gated or areolated appearance. Surface between the rugae deeply punctured. 
Metopidiimi more vertical than in concava, the percurrent carina more ele- 
vated and the dorsal sinus somewhat deeper. Apical margin of the head dis- 
tinctly sinuated next the eye, then very obtusely arcuated about the apex. 
Color blackish or fuscous, speckled or blotched more or less with testaceous on 
the head and front of the pronotum as far as the dorsal sinus; and with a tri- 
angular whitish testaceous spot on the apical fourth of the lateral margin, 
which may be prolonged somewhat along the margin anteriorly and indis- 
tinctly across the disk as an incomplete subapical vitta. Venter, tibiae and 
tarsi pallid, the disk of the ventral segments more or less black. 

Length, 4 mm. 

Distribution. Van Duzee reports this species from New Jersey, 
Pennsylvania, North Carolina and Missouri. There is a specimen 
in the Snow collection from Kansas City, Mo., and it has been taken 
in Riley county. 

Hosts. Unknown. 



94 The University Science Bulletin. 



PLATE II. 

1. Styles and coqgctive of Ceresa bubalus. 

2. ffidagus of Ceresa bubalus. 

3. Lateral aspect of Ceresa taurina. 

4. Genitalia of Ceresa taurina. 

5. Styles and connective of Ceresa bubalus. 

6. ffidagus of Ceresa bubalus. 

7. Genitalia of Ceresa bubalus. 

8. Lateral aspect of Ceresa bubalus. 



Lawson: Membracid^ of Kansas. 



95 



PLATE 11. 




96 The University Science Bulletin. 



PLATE III. 

1. Genitalia of Ceresa dicer os. 

2. ffidagus of Ceresa diceros. 

3. Styles and connective of Ceresa palnieri. 

4. ffidagus of Ceresa palmeri. 

5. Styles and connective of Stictocephala jestina. 

6. Lateral aspect of Stictocephala jestina. 

7. ffidagus of Ceresa borealis. 

8. Genitalia of Ceresa borealis. 



Lawson: Membracid.e of Kansas. 



97 



PLATE IIT. 




98 The University Science Bulletin. 



PLATE IV. 

1. Genitalia of Stictocephala inermis. 

2. CEdagus of Stictocephala inermis. 

3. Genitalia of Stictocephala lutea. 

4. ffidagus of Stictocephala lutea. 

5. Genitalia of Stictocephala inermis. 

6. Qildagus of Stictocephala inermis. 

7. Lateral aspect of Stictocephala inermis. 

8. Styles and connective of Stictocephala inermis. 



Lawson: Membracid.e of Kansas. 



99 



PLATE IV. 











100 The University Science Bulletin. 



PLATE V. 

1. Lateral aspect of genitalia of Thelia bimaculata. 

2. Dorsal aspect of genitalia of Thelia bimaculata. 

3. Dorsal aspect of genitalia of Micrutalis calva. 

4. Lateral aspect of genitalia of Micrutalis calva. 

5. Dorsal aspect of genitalia of Acutalis tartarea. 

6. Lateral aspect of genitalia of Acutalis tartarea. 

7. Lateral aspect of Telamona pyramidata. 

8. Styles and connective of Telamona pyramidata. 



Lawson: Membracid.e of Kansas. 



101 



PLATE V. 




102 The University Science Bulletin. 



PLATE VI. 

1. Lateral aspect of genitalia of Ophiderma salamandra. 

2. Dorsal aspect of genitalia of Ophiderma salamandra. 

3. Lateral aspect of genitalia of Publilia modesta. 

4. Dorsal aspect of genitalia of Publilia modesta. 

5. Dorsal aspect of genitalia of Entylia concisa. 

6. Lateral aspect of genitalia of Entylia ccmcisa. 

7. Dorsal aspect of genitalia of Vanduzea triguttata. 

8. Lateral aspect of genitalia of Vanduzea triguttata. 

9. Lateral aspect of genitalia of Campylenchia latipes. 

10. Ventral aspect of genitalia of Campylenchia latipes. 

11. Dorsal aspect of genitalia of Enchenopa binotata. 

12. Lateral aspect of genitalia of Enchenopa binotata. 



Lawson: Membracid.e of Kansas. 



103 



PLATE VI. 




104 The University Science Bulletin. 



PLATE VII. 

1. Lateral aspect of genitalia of Smilia camelus. 

2. Dorsal aspect of genitalia of Smilia camelus. 

3. Dorsal aspect of genitalia of Cyrtolobus querci. 

4. Lateral aspect of genitalia of Cyrtolobus querci. 

5. Dorsal aspect of genitalia of Cyrtolobus vau. 

6. Lateral aspect of genitalia of Cyrtolobus vau. 

7. Dorsal aspect of genitalia of Telamona viridia. 

8. Lateral aspect of genitalia of Telamona viridia. 



Lawson: Membracid.e of Kansas. 



105 



PLATE VII. 




INDEX. 

V.\C, K 

ac'uiniiuitus, tJki.s.suiiutus 67 

Acutalis ()2 

tartarea 62 

albescens, Ceresa. 52 

Archasia 78 

l)elfragei 78 

galeata 79 

Key to species 78 

Atynina 85 

belfragei, Archasia 78 

bimaculata, Thelia 66 

binotata, Enchenopa 49 

l)()realis, Ceresa 54 

brevicornis, Ceresa 55 

bubalus, Ceresa 57 

calva, JMicrutalis 63 

camelus, Smilia 80 

Campylenchia 47 

latipes 47 

carvff, Microcentrvis . . .' 45 

Carynota 64 

mera 64 

celsus, Cyrtoloyjus 81 

Centrotinae 45 

Ceresa 51 

albescens. . 52 

liorealis 54 

l)revicornis 55 

bubalus 57 

diceros 51 

Key to species 51 

palmeri 53 

taurina 56 

cinereus, Cyrt<)Iol)Us 84 

collina, Telamona 75 

concava, Pulililia 92 

concisa, Entylia 90 

cristata, Heliria 68 

Cyrt Globus 80 

celsus 81 

cinereus 84 

fenestratus 82 

f uliginosus 82 

griseus 83 

Key to subgenera 81 

muticus 86 

querr'i 85 

(107) 



108 The University Science Bulletin. 



PAGE 



Subgenus 81 

Key to species 81 

tuberosus 83 

vau 84 

decorata. Telamona 72 

diceros, Ceresa " 51 

Distribution 32 

Economic importance 43 

Enchenopa 48 

binotata 49 

Entylia 90 

eoncisa 90 

extrema, Telamona 74 

fenestratus, Cyrtolobus 82 

festina, Stictocephala 61 

flava, Ophiderma 89 

flaviguttula, Ophiderma 88 

f uliginosus, Cyrtolobus 82 

_galeata, Archasia 79 

Genitalia, male 38 

Glossonotus 67 

acuminatus 67 

griseus, Cyrtolobus 83 

Heliria 68 

cristata 68 

Key to species 68 

scalaris ._^ 68 

inermis, Stictocephala 59 

latipes, Campylenchia 47 

Life history 41 

List of species ' . . . . 44 

lugubris, Telamona 72 

lutea, Stictocephala ; 60 

Membracins 47 

Key to genera 47 

mera, Carynota 64 

Microcentrus 45 

carya3 45 

Micrutalis 63 

calva 63 

modesta, Pul)lilia 92 

modesta, Telamonanthe 78 

muticus, Cyrtolobus 86 

obsoleta, Telamona 71 

Ophiderma 87 

flava 89 

flaviguttula 88 

Key to species 87 

salamandra 87 



Lawson: Membracid^ of Kansas. 109 

PAGE 

palineri. Ceresa 53 

Phylogeny 41 

Puialilia 91 

concava 92 

Key to species 91 

modesta ' 92 

reticulata 93 

pyramidata, Tclainona 70 

querci, Cyrtolobus 85 

querci, Telamona 73 

reticulata, Publilia 93 

rileyi, Telamonanthe 77 

salamandra, Ophiderma 87 

scalaris, Heliria 68 

Smilia 79 

cainelus 80 

Smiliinae 50 

Key to genera 50 

Stictocephala 59 

festina 61 

inermis 59 

Key to species 58 

lutea ■ 60 

Structural characteristics 36 

Subfamilies, Key to .• 45 

tartarea, Acutalis 62 

taurina, Ceresa 56 

Telamona 69 

collina 75 

decorata 72 

extrema 74 

Key to species 69 

lugubris 72 

obsoleta 71 

pyramidata 70 

querci . 73 

unicolor 74 

viridia 71 

Telamonanthe 77 

Key to species 77 

modesta 78 

rileyi 77 

Thelia 65 

biniaculata 66 

Key to species 65 

uhleri 65 

triguttata, Vanduzea 89 

tuberosus, Cyrtolobus 83 

uhleri, Thelia 65 



110 The University Science Billetin. 

PAOE 

unicolor, Telamona 74 

Vancluzea 89 

triguttata 89 

vail, Cyrtolobus , . 84 

viridia, Telamona 71 

Xantholobus 86 



THE 

KANSAS UNIVEKSITY 

Science Bulletin 



Vol. XIV, No. 4— October, 1922. 

(Whole Series, Vol. XXIX, No. 4.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

The Genus Acinopterus (Homopter.\) P. B. Lawson. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 4. 



The Genus Acinopterus (Homoptera, Cicadellidae) , 

BY PAUL B. LAWSON, 
Professor of Entomology, University of Kansas. 



INTRODUCTION. 

THE genus Acinoptenis was erected by Van Duzee in 1892 to 
accommodate the species Acinoptenis acuminatus. Later, in 
1895, Baker described a single specimen as Phlepsius inornatus, a 
species which Van Duzee in his catalogue lists as Acinopterus acu- 
minatus var. inornatus. Finally, in 1903, Ball described three varie- 
ties of the typical species of the genus, the first of which, variety 
variegatus, Van Duzee made a synonym of Baker's variety inorna- 
tus. Thus up to the time that the writer undertook this study, one 
species and three varieties were recognized as comprising the mem- 
bership of this genus. The results of the study show that we now 
have at least eight species and one variety. One or two other species 
are probably represented in the material at hand, but because of the 
scarcity of specimens it was not thought best to describe them as new 
species. 

The writer is indebted to the following workers for very kindly 
loaning him the material studied : Mr. E. L. Dickerson, Mr. George 
G. Ainslie, Mr. F. H. Lathrop, Prof. J. G. Sanders, Dr. Dwight 
M. De Long, Mr. C. E. Olson, Mr. Edmund H. Gibson, Prof. H. 0. 
Osborn, and Dr. E. D. Ball. The work done is based for the most 
part on the many specimens from the collection of Doctor Ball, who 
not only loaned him all his own material, but also the types in the 
collection of the Iowa State Agricultural College, and in addition 
was a great help in making suggestions and giving information as 
to distribution and host plants. To Mr. E. H. Gibson and Mr. W. 
L. McAfee the writer owes the privilege of examining the collection 

(113) 



114 The University Science Bulletin. 

of the National Museum, including the type of Baker's inornatus. 
Through the kindness of Dr. J. Chester Bradley, the material at 
Cornell University was studied, and Prof. C. P. Gillette kindly per- 
mitted the examination of the collection of the Colorado Agricul- 
tural College. 

DISTRIBUTION. 

As far as our present knowledge goes, the members of this genus 
have hitherto been taken only in the United States and Mexico, a 
single specimen at least having been collected as far south as Yuca- 
tan. In our country it is for the most part southern and western in 
its distribution. This, along with its occurrence in Mexico, would 
indicate its probable presence throughout Central America and per- 
haps in the northern portion of South America. For this last state- 
ment, however, there is no certain data. The genus is, therefore, 
both Nearctic and Neotropical. 

The following states have yielded specimens of this genus: New 
Jersey, Maryland, District of Columbia, North Carolina, South 
Carolina, Virginia, West Virginia, Florida, Alabama, Mississippi, 
Tennessee, Arkansas, Missouri, Kansas, Oklahoma, Colorado, Utah, 
Washington, California, Arizona, Texas and Georgia. 

HOST PLANTS. 

Very little information is available as to the specific plants on 
which the members of this genus feed. Most of the specimens 
collected were taken while sweeping grasses or weeds. De Long 
reports Acinopterus acuminatus as abundant on grasses in Tennes- 
see. Ainslie took specimens from blue grass. Doctor Ball believes 
that A viridis var. variegatus feeds on wild geranium. A single 
specimen of A. angulatus was swept from Sphoeralcea angustifolia. 
A label on a specimen of A. acuminatus from Oklahoma states that 
it was taken from Amphiachyris, while another specimen from 
Rosser, Tex., was taken from Tetraneuris. Evidently most of the 
members of the genus are either grass feeders or else have as their 
hosts various weeds found in grassland. Much more careful collect- 
ing will need to be done before we can be certain of the specific 

hosts. 

DESCRIPTION OF THE GENUS. 

The following is Van Duzee's description of the genus: 

General appearance of Allygus, but with the elytra strongly narrowed 
posteriorly, and the tip acute. 

Head narrower than the pronotum, rounded, or somewhat produced before, 



Lawson: Genus Acinopterus. 115 

with the apex subacute, hind edge broadly concave. Vertex rather short, 
sloping, convex or more or less impressed behind the apex, surface punctured, 
the anterior submargin obscurely transversely rugose, passage to the front 
roimded. Front rather broad, at the base slightly encroaching upon the apex 
of the vertex, suddenh^ narrowed at tip. Clypeus much widened apically. 
Lorae large. Cheeks wide. Pronotum broad and rather short, anterior edge 
broadly arcuate, posterior nearly straight; sides long, oblique, carinated; 
lateral angles prominent, latero-posterior rounded. Scutcllum rather small. 
Elytra narrow, lanceolate at apex, the tip acute, appendix wanting; sutural 
edge straight to the extreme tip; costal and apical margins continuous; apical 
areoles five, inner small, oblique, second largest, reaching the extreme tip; 
third and fourth small; fifth, or stigmatal, long and usually crossed by one or 
two transverse veinlets; first and second sectors united by but one transverse 
nervure, but there are usually three or four connecting the outer claval nervure 
with the claval suture; all the nervures strong; costa feebly convex. Super- 
numerary cell of the wings present. Otherwise as in Athysanus and Allygus. 

The above description needs modification to enable it to include 
the several species in which the tegmina, while visibly narrowed 
apically, do not end in an acute tip, the sutural margin in these 
cases not continuing on straight to the extreme apex, but meeting 
the costal margin to form a rather broadly rounded tip. The writer, 
therefore, proposes the following generic description: 

Head usually narrower than pronotum, rarely as wide. Vertex rounded or 
distincth' angulate, usually impressed behind the apex, sloping, broadly round- 
ing with the front. Front broad basally, much narrowed apically. Clypeus 
widened apically. Lorse large, nearly reaching margin of the wide gense ven- 
trally. Pronotum wide, over twice as wide as long, anterior and posterior 
margins nearly parallel, latter usually more nearly straight; lateral and 
humeral margins distinct; disc transversely wrinkled. Scutellum finely granu- 
lar and with distinct transverse impressed line. Tegmina moderately long, apex 
alwaj's narrowed, sometimes to an acute tip. Venation distinct. Of the five 
apical cells M4 is the largest, but it, along with Cui and Ro, is frequently 
divided by cross-veins, R2 being so regularly divided that probably both Ri 
and R2 are present. Cell 2d M is not divided by a cross-vein. Veins 1st A and 
2d A almost always connected by one or more cross-veins. Valve of male 
always hidden under last ventral segment; plates usually long but never 
equaling pygofers. Female, last ventral segment large, always bearing a median 
notch. 



116 The University Science Bulletin. 

SYSTEMATIC TREATMENT OF THE SPECIES. 

KEY TO SPECIES. 

A. Apex of tegmina strongly acute; costal margin straight clear to tip. 
B. Brownish or greyish species. 

C. Darker and larger species. 

D. Male plates long and narrow, parallel-margined, apices 
rounded; last ventral segment of female without lateral 
angles. acuminatus Van D. 

DD. Male plates broad and shorter, apices distinctly diver- 
gent; last ventral segment of female with distinct lateral 
angles. angulatus n. sp. 

CC. Lighter and more slender species. inornatus (Bak.). 

BB. Greenish species. productus n. sp. 

AA. Apex of tegmina rounded; costal margin not running straight clear to tip. 
B. Species brownish or greyish, not distinctly green. 

C. Species large; last ventral segment of female strongly pro- 
duced medially. brunneus Ball. 
CC. Species smaller; last ventral segment of female normally 
produced. 

D. Species with elytra distinctly variegated. 

viridis var. variegatus Ball. 

DD. Species with elytra not variegated pallidus n. sp. 
BB. Greenish species. 

C. Species larger; plates of male tapering and distinctly divergent 
apically. viridis Ball. 

CC. Species smaller; plates of male parallel-margined and but 
slightly divergent apically. obtutus n. sp. 

Acinopterus acuminatus Van D. 

(PI. VIII, fig. 1; pi. IX, fig. 1; pi. X, fig. 1; pi. XI, fig. 1; pi. XII, figs. 1, 2.) 
Acinopterus acuminatus Van D., Psyche, vi, p. 308, 1892. 
Acinopterus acuminatus G. & B., Heniip. Colo., p. 94, 1895. 

Acinoptervs acuminatus Van D., Bui. Buf. Soc. Nat. Sci., viii. No. 5, p. 69, 1907; ix, p. 
225, 1909. 

Acinopterus acuminatus Osb., Ohio Nat., ix, p. 466, 1909. 
Acinopterus acuminatus Smith, Cat. Ins. N. J., edn. 3, p. 105, 1910. 
Acinopterus acuminatus Barb., Bui. Am. Mus. Nat. Hist., xxxiii, p. 534, 1914. 
Acinopterus acuminatus Van D., Trans. San Diego Soc. Nat. Hist., ii, p. 54, 1914. 
Acinopterus acuminatus Mete, Jl. Elisha Mitchell Sci. Soc, xx.xi, p. 23, 1915. 
Acinopterus acuminatus De L., Tenn. St. Bd. Ent., Bui. 17, p. 89, 1916. 
Acinopterus acujninatus Van D., Cat. Hemip. N. A., p. 675, 1917. 
Acinopterus acuminatus Lathr., S. C. Agr. Exp. Sta., Bui. 199, p. 102, 1919. 
Acinopterus acuminatus Laws., Kans. Univ. Sci. Bui., xii, p. 207, 1920. 

The following is the original description of this species: 

Fulvous brown tinged with dull green or yellowish, elytral nervures pale, 
brown-margined. Length, 5-6% mm. 

Head pale. Front with about eight brown arcs, more or less distinct. 
Pronotum feebly calloused on the anterior margin, with a few shallow im- 
pressions arranged parallel to the edge, more obvious in the males; lateral 
margin as long as the latero-posterior, acutely carinated; disc posteriorly ob- 
scurely wrinkled. Basal angles of the scutellum with a brownish triangular 
spot more or less apparent. Pectoral pieces usually more or less invaded with 
blackish, sometimes pale and immaculate. Legs pale, or suffused with 



Law son: Genus Acinopterus. 117 

sanguineous. Abdomen pale, frequently black above, excepting the broad 
lateral margins; infuscated on the basal and apical segments of the venter in 
the females; slightly suffused with a pale median line in the males. Elytra 
pale fulvous, frequently whitish hyaline on the disc of the costal and some of 
the discal areoles of the corium, and on the inner margin of the clavus, the 
extreme apex clouded with smoky or even blackish; nervures pale, edged 
with brownish, the marginal with a fuscous interruption at tip; claval suture 
brown. Wings smok}^ iridescent, nervures fuscous. 

Genitalia. Male: Valve wanting. Plates long and narrow, a little longer 
than the last ventral segment, about one-third wider at base than at 
their obtusely lanceolate, divergent tips. Pygofers twice the length of the 
plates, narrowed and obtusely pointed at apex, armed beyond the plates with 
numerous stout spines. Female: Last ventral segment rather long, hind edge 
with a shallow median notch, either side of which is a broadly roimded lobe, 
retreating at the outer angles. Pygofers rather broad, their subacute apex 
moderately exceeded by the oviduct. 

Described from 5 males, 3 females. Maryland, September 29 and August 4, 
on pines (Uhler) ; North Carolina (Osborn) ; New Jersey (Uhler) ; mountains 
of northwest Colorado (Gillette); California (Coquillett). 

In a male from California the lower surface of the femora is black. A 
female from North Carolina has the disc of the elytra white-pruinose, and all 
the specimens exhibit considerable variation in the extent of the black 
markings. 

The writer gives the following description : 

A large, rather robust, brownish species, ranging from a greyish-brown to a 
usually dark-brown color. Length, 5 to 7 mm. 

Form. Head distinctly narrower than the pronotum. Vertex usually 
distinctly produced medially, about half longer at the middle than next the 
eyes and about twice as broad as long. Front broad, lorae large, cheeks wide, 
clypeus widened apically. Pronotum over twice as wide as long, anterior 
margin usually a little more curved than posterior, the lateral and humeral 
margins subequal, the disc transversely wrinkled. Scutellum large, the surface 
granular. Tegmina moderately long, the costal margin running straight 
clear to the apex, forming an acute tip. 

Color. Vertex, pronotum and scutellum brownish to olive green, the 
scutellum with basal angles and three longitudinal lines, light. Tegmina usually 
shining dark brown, sometimes lighter. When dark brown the nervures are 
lighter and some of the cells, especially along the costa and on the clavus, are 
subhyaline or greenish. Light specimens have the viens, especially apically, 
bordered with brown. Face olive green, unmarked, or with faint arcs on the 
front. Below olive green, marked more or Ipss with dark brown or black, 
especially on the thorax, coxse and tergites of the abdomen. 

External genitalia. Female: Last ventral segment twice as long as preced- 
ing, broad basally, lateral margins rounding to slightly notched and produced 
posterior margin ; pygofers bearing a few scattered spines and slightly exceeded 
by ovipositor. Male: Valve not visible, plates long and narrow, parallel- 
margined, obtuse apices somewhat divergent and greatly exceeded by spiny 
P3'-gofers. 



118 The University Science Bulletin. 

Internal male genitalia. Styles with margins of anterior half sinuately 
tapering, distal half stout and strongly curved; the large, club-shaped and 
coarsely granular apices strongly diverging. Connective small, heart-shaped, 
with the excision wide and the apex broadly rounding. CEdagus broad basally, 
narrowing to the middle, bearing two small basal processes and a larger apical 
one near the tip of which, on the ventral surface, is the fimbriated opening of 
the penis. 

Distribution. This species is largely southern and eastern in its 
distribution. The many specimens examined by the writer, in addi- 
tion to those mentioned by Van Duzee, are distributed as follows: 
Charter Oak, Pa. (J. N. KnuU) ; Pt. Royal, Harrisburg, Rockville, 
Pa. (J. G. Sanders) ; Lakehurst, N. J. (J. B. Weiss) ; Great Falls, 
Md., Berkeley, W. Va., Ft. Royal, Va., Washington, D. C. (Heide- 
mann) ; Orangeburg, S. C. (F. H. Lathrop) ; Kansas City, Mo. (F. 
Rogers) ; Bisc Bay, Jacksonville, Fla., Gainesville, Fla. (C. J. 
Drake) ; Ardmore, Okla. (F. C. Bishopp) ; Jacksonville, Tex. (W. D. 
Pierce) ; Boerne, Tex. (F. C. Pratt) ; Victoria, Rosser, Tex. (J. D. 
Mitchell) ; Kushla, Ala. (A. H. Sturtevant) ; Alabama, Florida, 
Mexico (C. F. Baker) ; Knoxville, Nashville, Tenn. (W. B. Cart- 
wright, C. C. Hill) ; Nashville, Covey Spring, Chattanooga, Tenn. 
(Geo. G. Ainslie) ; Colliersville, Clarksville, Paris, Lexington, Tenn. 
(Dwight M. De Long) ; Agricultural College, Mississippi (H. E. 
Weed) ; Cherokee, Bourbon and Miami counties, Kansas (R. H. 
Beamer) ; Virginia, labeled Jassits tructilis, (Uhler) ; Spring Creek, 
Okefinokee Swamp, Bainbridge, Ga. (J. C. Bradley) ; Little Rock, 
Ark.; Capa, S. D. 

E. L. Dickerson reports what are presumably this species from 
Cologne, Lakehurst and Egg Harbor, N. J. 

Remarks. The writer has examined the three type specimens, one 
male and two females, of Adnopterus acuminatus from the collec- 
tion of the Iowa State College, and the female type from the Cornell 
LTniversity collection. The two female specimens from Maryland 
and Virginia are clearly of a different species from the male and fe- 
male from California, as shown by a comparison of the last ventral 
segment of the two females. It seems evident, though, that the ma- 
jority of the eight specimens from which Van Duzee described the 
species were from the East or Southeast, and that he evidently meant 
to describe a species with such a distribution. Accordingly these 
eastern females are retained as types of this species, while the two 
California specimens, along with a large amount of western material, 
are placed in the following species, which not only is clearly western 
in its distribution, but is decidedly different as to the genitalia of 
both males and females from the above species. 



Lawson: Genus AciNOPTERrs. 119 

Acinopterus angulatus n. sp. 

(PI. VIII, fig. 8; pi. IX, fig. 3; pi. X, fig. 2; pi. XI, fig. 4; pi. XII, figs. T), 6.) 

A smaller species than the preceding, vaiying from light" to dark brown in 
color, but uniformly lighter than acuminatus and lacking its olive-green tinge. 
Length, 5 to 6.25 mm. 

Form. Head broad, scarcely narrower than prothorax. Vertex broad and 
short, over twice as wide as long, about one-third longer at middle than next 
the eyes, anterior margin rounded or slightly angulate, sloping, and with an 
impressed line behind the apex. Front broad and short, lorse, clypeus and 
gense characteristic of the genus. Pronotum over twice as wide as long, an- 
terior margin broadly convex, posterior margin shallowdy concave, lateral and 
humeral margins distinct, the disc transversely wrinkled. Scutellum finely 
granulated, the transverse impressed line curved and distinct. Tegmina with 
sutural margin rimning straight clear to the tip, forming an acute apex. 
Venation distinct, with from one to several cross-veins between the first and 
second anal veins. 

Color. Vertex, pronotum and scutellum browTi or yellowish brown. Scu- 
tellum with basal angles and three longitudinal lines, light. Elytra brown, the 
veins margined with darker brown, so that many of the cells, especially along 
the costa and apicallj^ appear light. Face browTi; front with eight pairs of 
dark-brown lines, wliich are longest above and shortest below, leaving the mid- 
dle portion unmarked. Below browm, but with parts of thorax, the coxae and 
femora of the meso- and metathoracic legs and the dorsum of the abdomen 
usuall}- black or dark brown. 

External genitalia. Female: Last ventral segment twice as long as pre- 
ceding, the posterior margin varying from slightly concave to slightly convex, 
but always with a shght median notch and foiTning very distinct lateral angles 
with the long lateral margins. Pygofers moderately wide, sparsely spined, ex- 
ceeded shghtly bj^ the ovipositor. Male: Last ventral segment long, hiding 
the valve. Plates broad, about the length of the last ventral segment, only 
slight h' narrowing to the obtuse tips which are divergent medially, but have 
distinct lateral angles. Pygofers with a few stout spines and exceeding the 
plates by about two-thirds the length of the latter. 

Internal male genitalia. Styles with rather small and pointed anterior ends, 
widest at point of the distinct process to the connective, then strongly curved 
to the very wide apices, which are quite granulated and clearly concave be- 
tween the outer angle and the larger, more produced inner apex. The outer 
margin bears a few small spines. Connective heart-shaped, the apex quite 
broad. (Edagus of the pattern characteristic of the genus, the basal processes 
well developed and toothed, the terminal process veiy long, clearly showing 
the penis, which opens at the fimbriate extreme apex of the process. 

Distribution. With one exception, a specimen from Washington, 
D. C, all the material at hand came from the South and West. A 
single specimen was taken in Yucatan, many specimens coming 
from other parts of Mexico. We are evidently safe in calling it a 
southwestern species, which may possibly extend its range into the 
southeastern states. The following are the locality records of the 



120 The University Science Bulletin. 

material studied: Riverside, Chino, San Diego, Ontario, Visalia, 
Tia Juana, Oroville, Caliente (Ball) ; Hamilton (no collector's 
name) ; Los Angeles (Coquillett and A. Koebele) ; Whittier, Brawley 
(H. 0. Osborn) ; all from California, along with a number of speci- 
mens taken by C. F. Baker; Vera Cruz (no collector's name) along 
with two specimens taken by Gillette and a series by Baker, all from 
Mexico; Victoria (J. D. Mitchell), Orizaba (H. Osborn), College 
Station (no collector's name), Brewster county (Mitchell and Cush- 
man), all from Texas. Also a single specimen taken at Progresso, 
Yucatan, by Frederick Knab, and one taken at Washington, D. C. 

Holotype, male, Ontario, Cal. Collection of E. D. Ball. 

Allotype, female, Ontario, Cal. Snow collection, University of 
Kansas. 

Paratypes, male, Visalia, Cal., and female, Chino, Cal., in col- 
lection of E. D. Ball; male, Riverside, Cal., and female, Chino, Cal.. 
in Snow collection; male, Whittier, Cal., and female, Brawley, Cal., 
in collection of H. 0. Osborn. 

Acinopterus inornatus (Bak.) 

(PI. VIII, figs. .3, i [type], ,'5; pi. IX, figs. 8, 9; pi. X. fig. (1; pi. XI, fig. 7; 

pi. XII, figs. 11, 12.) 

Phlepsius inornatus Bak., Psyche, vii, Suppl. i, p. 13, 1895. 

Acinopterus acuminatus var. inornatus A'an D., Cat. Heniip. N. A., p. 675, 1917. 

The following is the original description. 

Phlepsius inornatus, n. sp. Differing from all other species of the genus 
in being entirely destitute of elytral reticulations or other markings. Length 
of male, 6 mm. 

Male: Head narrower than the pronotum. Face a twelfth wider than long; 
clypeus one-half longer than broad, somewhat constricted before the base, basal 
suture strongly curved, apex slightly concave; lorae as long and two-thirds as 
broad as clypeus; margin of gense rather slightly incur\-ed below the eye, below 
this strongly conve.x, thence slightly incurved to tip of clypeus. Front an 
eleventh longer than broad, somewhat less than twice the length of the 
clypeus, broad below, the sides very slightly incurved at the antennse. Disc 
of the vertex flat, length at middle once and a half that next the eye, width 
between the eyes once and a half the length. Width of the pronotum two and 
a third times the length, the length about once and two-thirds that of the 
vertex, curvature nearly two-fifths of the length, posteriorly irregularly trans- 
versely wrinkled. Scutellum and elytral venation normal. Plate not visible, 
valves two and a half times longer than broad at base, slightly narrower at 
apex, blunt at tips, without hairs. Pygofers one-half longer than valves, 
pointed at tips, their whole outline subtriangular, provided on disc of lower 
surface with several rather long whitish spines arranged in a single longitudinal 
row. 

Color very pale yellowish, deeper on the abdomen. Pronotum with five 
very indistinct longitudinal whitish bands. Elytra translucent, pale milky 



Lawson: Genus Acinopterus. 121 

white, with indistinct smoky clouds on the discs of some of the apical and 
anteapical areoles. Veins white, claval suture brownish. Face and legs tinged 
with greenish, some of the white tibial spines brown tipped. Tarsal joints at 
apico;* dark. Dorsal abdominal segments, fexcept lateral and apical margins, 
blackish. 

Described from a single male taken at San Augustine (Ckll. 2140). In form 
this insect very closely resembles P. superbus and in structure is strictly con- 
generic with it. It differs very widely, however (and this is a generic differ- 
ence according to Van Duzee's sj'noptic table of the genera), in that it does not 
possess the elytral reticulations or other markings so characteristic of the 
genus. On a very superficial examination it might be taken for a Chlorotettix, 
but its general form, stronger build, and lengthened vertex are strictly Phlep- 
siid. 

The writer gives the following description: 

A rather slender and light-colored species which sometimes may be rather 
dark. Length, 5 to 6.5 mm. 

Form. Head distinctly narrower than the pronotum. Vertex varying in 
length, but usually about twice as wide as long and half longer at the middle 
than next the eye; disc sloping and with the impressed line behind the apex. 
Face as in the other members of the genus. Pronotum over twice as wide 
as long, the anterior margin more strongly curved than the posterior, the 
lateral and humeral margins about equal. Scutellum of average size, finely 
granulated, and with distinct transverse impressed line. Tegmina long and 
narrow, sutural margin extending straight clear to tip, forming an acute 
apex; venation usually distinct, though sometimes rather weak, with from one 
to several cross-veins between the first and second anal veins. 

Color. Yellowish or yellowish grey. Pronotum with five pale and some- 
times indistinct lines. The tegmina vary considerably, being sometimes 
almost colorless till near the tip, while in others the veins are margined lightly 
with brown, especially at the margins, but in all cases some of the apical cells 
are more or less darkened. The darkened tips of the veins along the sutural 
and costal margins sometimes give the elytra a variegated appearance. Be- 
neath this species is usually light except for the darkened doreum of the 
abdomen. 

External genitalia. Female: Last ventral segment over twice as long as the 
preceding, posterior margin di.stinctly but roundingly produced medially, but 
with the small median notch characteristic of the genus. Pygofers moderately 
wide, sparsely spined, and slightly exceeded by the ovipositor. Male: Last 
ventral segment longer than the preceding, hiding the valve. Plates long and 
fingerlike, about the length of the last ventral segment, shghtly narrowed to 
the somewhat diverging but rounded tips. Pj'gofers bearing a few stout spines 
and exceeding the plates by about two-thirds the length of the latter. 

Internal male genitalia. Styles wide basally and with strong process to con- 
nective; apically strongly curv^ed and clublike, the apical portion of the club 
with distinct and large granulations, giving it a rough appearance. Connective 
heart-shaped, the excision wide and shallow, the apex rounded, ffidagus rather 
small but stout, the upper part much like an inverted boot, the heel distinctly 
cleft ; the paired basal processes about half the length of the apical process, 
their lower edges serrate; the terminal process stout, the penis opening at the 
fimbriate tip. 



122 The University Science Bulletin. 

Distribution. This species is evidently a southwestern form, for 
all the material at hand was taken in this region. The following 
are the locality records of the twenty-two specimens examined: 
Type specimens from San Augustine, N. M. (C. F. Baker) ; On- 
tario, Riverside, Cal. (E. D. Ball) ; Santa Rita mountains (F. H. 
Snow), Galiuro mountains (H. G. Hubbard), Phoenix (E. D. Ball), 
Sabino Canyon, St. Catalina mountains (E. L. Dickerson) , all from 
Arizona; Brewster county, Texas (Mitchell and Cushman). 

Remarks. As far as the writer knows, this species has been known 
hitherto only from the type. In the material gathered for the study 
of the genus he found a number of similar specimens, which, while 
differing in some ways, particularly in the length of the vertex and 
the extent of the elytral markings, are yet thought to be representa- 
tives of this species, for a careful study of the genitalia of several 
males revealed no differences, although the vertices of the specimens 
were quite unlike. Also the specimens show a complete range in 
color from that of the very light type specimen to forms that are 
distinctly brownish. While frankly having some doubt as to the 
specific identity of all the material named thus, the writer feels it 
better to call them all the same species rather than to describe new 
species on insufficient material. 

Acinopterus productus n. sp. 

(PI. VIII, fig. G; pi. IX, fig. 2; pi. X, fig. 3; pi. XI, fig. 3; pi. XII, figs. 13, 14.) 

A distinctly greenish species, differing from other green forms by the acute 
apex of the tegmina and the produced vertex. 

Form. Head distinctly narrower than pronotum. Vertex less than twice as 
wide as long, at least half longer at the middle than next the eye, the apex 
rounded and with an impressed line parallel with the margin. Face character- 
istic of the genus. Pronotum short, well over twice as wide as long, the ante- 
rior margin more strongly curved than the posterior, the lateral and humeral 
margins about equal, the disc transversely wrinkled. Scutellum with the usual 
granular surface 8,nd impressed line. Tegmina long and rather narrow, the 
costal margin running straight clear to the tip, forming an acute apex, and 
with the nervures distinct, the fii-st and second anal veins usually united by 
several cross-veins. 

Color. The entire insect is green except for the darkened apices of the ely- 
tra and the yellowish or pinkish legs. The nervures stand out as a lighter 
green than the cells of the tegmina. 

External genitalia. Female: Last ventral segment over twice as long as 
the preceding, the medially produced posterior margin with the usual small 
notch, the lateral margins rounding with the posterior. Pygofers rather ro- 
bust, sparsely spined, and slightly exceeded by the ovipositor. Male: Valve 
hidden by the long last ventral segment. Plates longer than last ventral seg- 
ment, fairly wide basally, tapering to the divergent and rounded but com- 



Lawson: Genus Acinopterus. 123 

parativeh' narrow ajiex. Sparsely bristled pygofers exceeding i)]a1es by about 
two-thirds the length of the latter. 

Internal male, (icnitalia. Styles of usual form, widest at point of process to 
connective, apically club-shaped, the granulated and blunt apices slightly but 
clearly concave. Connective heart-shaped, the excision fairly deep, ffidagus 
with body as in other species, the basal processes small, the terminal process 
of medium diameter and length. 

Distribution. The eight specimens at hand when this species was 
described all came from California and Arizona. They were all 
taken by Dr. E. D. Ball. The California specimens are from Im- 
perial, Beaumont and Riverside, while the two Arizona specimens 
are from Phoenix. 

Holotype, female, Imperial, Cal.. in collection of Doctor Ball. 

Allotype, male, and paratype, female, both from Imperial, Cal., 
in the Snow collection, University of Kansas. 

Paratypes, male from Imperial and female from Beamnont, Cal., 
in collection of Doctor Ball. 

Acinopterus brunneus Ball. 

(PI. VIII, fig. 2; pi. IX, fig. 5; pi. X, fig. 7; pi. XI, fig. 6; pi. XII, figs. 9, 10.) 

Acinopterus acuminatus var. brunneus Ball, Can. Ent., xxxv, p. 231, 1903. 
Acinopterus acuminatus var. brunneus Van D., Cat. Hemip. N. A., p. 675, 1917. 

The following is the original description : 

^4. acuminatiifi, var. brunncuf;, n. var. Shghtly larger than the preceding 
variety. "\'ertex, pronotum and scutellum pale green, washed with cinnamon 
brown. Elytra pale cinnamon brown, slightly fuscous at tip. Whole insect 
with a slight tawny iridescence, below pale green. 

Described from three specimens from Rifle, Colo., taken by the author. 

• 
The writer adds the following description: 

A large brownish or greenish-brown species, about the largest member of 
the genus. Length, 5.5 to 6.75 mm. 

Form. Head about as wide as the pronotum. Vertex at least twice as wide 
as long, one-third longer at middle than next the eye, the anterior margin 
rounded, and with the characteristic depression behind the apex. Face with 
all the parts ver\' broad, the lorse nearly reaching the margin of the genae. 
Pronotum over twice as wide as long, the anterior margin more strongly cur\^ed 
than the posterior, lateral and humeral margins distinct and about equal, the 
disc transversely WTinkled. Scutellum as in other members of the genus. Teg- 
mina with rounded apex but more acute than in viridis, the venation often less 
distinct than in other species, and usually with several cross-veins between the 
first and second anal veins. 

Color. Vertex, pronotum and scutellum greenish-brown. Tegmina of same 
color or darker, the veins of the apical half often being margined with dark 
brown, giving the tip a darker appearance. Beneath the color is usually as 
above but sometimes the hind legs and abdomen have a reddish tinge. 



124 The University Science Bulletin. 

External genitalia. Female : Last ventral segment differing from that of 
any other member of the genus in being extremely produced medially, three 
times as long as the preceding segment, with the usual small apical excision, 
and the lateral margins sometimes slightly concave. The broad and spiny 
pygofers are slightly exceeded by the ovipositor. Male: Last ventral segment 
long, hiding the valve. Plates long and slender, longer than last ventral seg- 
ment, their sides straight to the somewhat narrowed but rounded apices. 
Pygofers long, exceeding plates by about the length of the latter, bearing the 
usual spines. 

Internal male genitalia. Styles stout, apical part nearly of same width 
throughout and bearing many granulations. Connective as broad apically as 
basally. CEdagus unhke anything in the genus and very characteristic, the 
basal processes short and not serrate, the distal process verj' long, the fimbriate 
opening of the penis extending back from the extreme tip for a considerable 
distance. In addition a pair of large ventral and lateral lobes is present that 
completely cover the basal processes, these lobes being serrate along their ven- 
tral margin. The entire dorsal part of the oedagus is also quite different from 
the corresponding part in the other species of the genus. 

Distribution. With the exception of a single specimen taken by 
Coquillett at Los Angeles, Cal., all the other fourteen specimens ex- 
amined by the writer are from Doctor Ball's collection. These were 
taken from the following localities: Rifle, Colo.; Pardman, Salt 
Lake City, Utah; Ravenna, Cabazon, Riverside, and Beamnont, Cal. 

Acinopterus pallidus n. sp. 

(PI. Vlll, fig. 10; pi. IX, fig. 4; pi. XI, fig. 2.) 

Closely related to the preceding species, but slightly smaller and paler. 
Length, 5.5 to 6 mm. 

Form. Head distinctly narrower than the pronotum. Vertex a little over 
twice as wide as long, not quite one-third longer at the middle than next the 
eye, the anterior margin evenly rounded and broadly rounding with the front, 
the depression back of the apex small. Face very broad, the front fully as 
broad basally as long, and the gense quite wide. Pronotum over twice as wide 
as long, anterior margin but slightly more curved than the posterior, lateral 
and humeral margins about equal, disc with transverse wrinkles. Scutelkun 
with usual granular surface and transverse impressed line. Tegmina moder- 
ately long, the apices more rounded than in preceding species, venation dis- 
tinct but not conspicuous, and with but one or two cross-veins between the 
first and second anal veins. 

Color. The entire insect, abov^e and below, of a pale greenish-yellow color. 
Apices of some of the veins along costal margin and at apex margined with 
black, making the tips of the tegmina appear slightly darkened. The tarsi 
tend to be brownish. 

External genitalia. Female : Last ventral segment produced medially, bear- 
ing a shallow median notch apically from which the margins round to the 
base. Pygofers large, sparsely spined, exceeded by the ovipositor. 

Distribution. Described from four specimens taken by Doctor 
Ball at Cabazon, Cal. 



Lawson: Genus Acinopterus. 125 

Holotype, female, and two paratypes, females, in collection of 
Doctor Ball. 

One paratype, female, in the Snow collection, University of 
Kansas. 

Remarks. This species stands between brunneus and viridis. 
From the former it differs in its smaller size, lighter color, and 
shorter female ventral segment, while from the latter it differs in 
being lighter, and in not having the tegminal veins green. 

Acinopterus viridis Ball. 

(PI. VIII, fig. 11; pi. IX, fig. 6; pi. X, fig. 4; pi. XI, fig. 5; pi. XII, figs. 7, 8.) 

Acinopterus acuminatus var. viridis Ball, Can. Ent., xxxv, p. 231, 1903. 
Acinopterus acuminatus var. viridis Van D., Cat. Hemip. N. A., p. 675, 1917. 
Acinopterus acrimirtatus var. viridis Laws., Kan. Univ. Sci. Bui., xii, p. 208, 1920. 

The following is the original description: 

A. acuminatus, var. viridis, n. var. Form and structure of the preceding 
nearly; slighth- smaller. Bright grass green both above and below. Eyes 
and extreme tip of elytra fuscous. 

Described from a number of specimens from southern Colorado and Ari- 
zona. This is the common form in southern Colorado, where it was taken by 
E. P. Van Duzee and the author. 

The following description is by the writer : 

A rather robust greenish species, with or without elytral markings. Length, 
5 to 6 mm. 

Form. Head distinctly narrower than the pronotum. Vertex about twice 
as wide as long, one-half longer at the middle than next the eye, anterior 
margin broadly rounded and with a depression just behind apex. Face with 
all the sclerites broad. Pronotum over twice as wide as long, anterior margin 
more strongly curved than the posterior, lateral and humeral margins about 
equal, the disc transversely wrinkled. Scutellum as in other members of the 
genus. Tegmina moderately long, the apex narrowed but rounded, with 
usually one or two or sometimes several cross-veins between the first and 
second anal veins and sometimes one or two between the second and third. 

Color. Vertex, pronotum and scutellum usuall}^ green, though sometimes 
distinctly j'ellowish. Tegmina green with the nervures darker green, the 
latter being sometimes not margined at all or bordered with brown till all 
the apices of the veins at the sutural margin, along the distal half of the 
costal margin, and at the apex, are definitely bordered, frequently giving the 
apex a darker appearance. Below the insect is also green, the tarsi tending 
to be brownish. 

External genitalia. Female : Last ventral segment over twice as long as 
the preceding, the lateral and posterior margins rounding to the produced apex 
which bears the usual small median notch. Pygofers stout, sparsely spined, 
exceeded by the ovipositor. Male: Last ventral segment long, hiding the 
valve. Plates long and slender, slightly longer than last ventral segment, the 
bases distinctlj' wider than the divergent apices. Pygofers broad, sparsely 
spined, exceeding the plates by about two-thirds the length of the latter. 

Internal male genitalia. Stjdes large, widest at point of process to connec- 



126 The University Science Bulletin. 

tive, distal portion strongly curved and then running straight to the expanded 
tips, which have the inner angles about right-angled, but the outer angles 
strongly produced, the distal margin between the two corners being slightly con- 
cave. The outer margins of the distal half are roughened and the characteris- 
tic granulations appear over the entire apical portion. Connective heart-shaped, 
the apex broadly rounded, ffidagus very characteristic of the species, having 
two pairs of basal processes, the upper ones shorter, the lower ones reaching 
nearly to the tip of the apical process, at the extreme fimbriate tip of which 
the penis opens. Both pairs of basal processes bear teeth along the margins. 

Distribution. All of the twenty-five specimens, except one from 
Colorado, one from Morton county, Kansas, and one from Ashfork, 
Ariz. (Barber and Schwarz), were sent the writer by Doctor Ball, 
who obtained them from the following localities: Soldier, Dixie, 
Richfield, Monroe, Moab, Utah; Fort Collins, Grand Junction, 
Delta, Dutch George or Poudre Canyon, Colo.; Coolidge, Kan.; 
Wenatchee, Wash. ; Phoenix, Ariz. There are specimens also in the 
collection of the Colorado Agricultural College from some of these 
localities. 

Remarks. The specimen from Moab, Utah, seems to be different 
from the other specimens of this species in that it is lighter in color, 
has a broader head, and longer and more pointed elytra. There be- 
ing only one specimen of its kind, however, the writer prefers to 
place it here to describing it as a new species from a single speci- 
men. 

Acinopterus viridis var. variegatus Ball. 

(PI. VIIl, fig. 7 ; pi. IX, fig. 7.) 

Acmopterus acuminatus var. variegatus Ball, Can. Ent., xxxv, p. 231, 1903. 
Acinopterus acuminatus var. inornatus Van. D., Cat. Hemip. N. A., p. 675, 1917. 

The following is the original description: 

Acinopterus acuminatus, var. variegatus, n. var. Form and structure of the 
species, but much lighter colored. Vertex, pronotum and scutellum inclined 
to be reddish, especially in the male. Elytra whitish pruinose, nervures green- 
ish, not margined, except towards apex and along sutural margin, three fuscous 
points along the suture, and sometimes one on the disc of each elytron. 

Described from twenty-four specimens from Colorado and Arizona. 

The following is the writer's description: 

The members of this variety are hke viridis except in color. 

Color. General color, brown. Vertex, pronotum and scutellum greyish or 
brownish, sometimes with a reddish tinge. Tegmina pale, but with nervures 
margined more or less throughout, especially along sutural and costal margins 
and apically, giving them a decidedly variegated appearance. 

Distribution. Eight specimens examined are from Doctor Ball's 
collection and were taken by him at Fort Collins and Denver, Colo. 



Lawson: Genus Acinopterus. 127 

In the collection of the Colorado Agricultural College are other 
specimens from the same localities. 

Remarks. The specimens at hand show gradual gradations into 
the variegated form of viridis, which, in its turn, goes by insensible 
gradations into the pure green form characteristic of the species. An 
examination of the male internal genitalia of typical variegatus and 
that of a variegated viridis showed no differences, and the gradual 
loss of the tegminal markings into the plain green form would seem 
to indicate the identity of these two green forms. 

Acinopterus obtutus n. sp. 

(PI. VIII, fig. 9; pi. X, fig. 5; pi. XII, fig.s. 3, 4.) 

A rather small green species with a relatively larger vertex than viridis. 
Length. 5.5 mm. 

Form. Head di.stinctl.>- narrower than pronotiim. Vertex large, about twice 
as wide as long, one-half longer at the middle than next the eye, a sHght de- 
pression just behind the broadly rounded apex which rounds very obtusely with 
the front. All the sclerites of the face rather broad. Pronotum over twice as 
wide as long, the anterior margin more curved than the posterior, the lateral 
and humeral margins about equal, the transverse wrinkles of the disc indis- 
tinct. Scutellum with the usual granulated surface and transverse impressed 
line. Tegmina with the tips broken off in all three of the specimens from 
which the species is described, but presumably rather rounded apically, judging 
from the material studied. Claval area with a few cross-veins between the 
first and second anal veins. 

Color. Vertex, pronotum and scutellum green, the scutellum with basal 
angles and three longitudinal lines, light. Tegmina milky green, the veins 
light or dark green, and margined more or less with brown. Below the entire 
insect is green. 

External genitalia. Male: Last ventral segment long, hiding the valve. 
Plates long and narrow, nearly parallel-margined to the somewhat divergent 
apices, which are exceeded by the bristly pygofers by about two-thirds the 
length of the former. 

Internal male genitalia. Styles of the usual shape, the granular apical por- 
tions quite expanded at the tip, which is either straight or slightly concave at 
the end. Connective nearly round, the basal excision rather deep. Qildagus 
differing from that of any member of the genus. The body is very wide 
ba.sally, the heel cleft, then strongly narrowed to the base of the procesvses, of 
which the basal pa'ir are short and slender while the distal one is quite long 
and slender, the penis opening at the extreme fimbriate tip. 

Distribution. The three male specimens from which this species 
is described are all from the collection of Mr. E. L. Dickerson and 
were taken in the Sabino canyon of the St, Catalina mountains of 
Arizona. 

Remarks. This species is decidedly smaller than the other green 
species, and the cedagus is so characteristic that on this structure 



128 The University Science Bulletin. 

alone the writer is confidently basing the distinctness of thefi6 spe- 
cies. Among the specimens of viridis from Colorado and Utah there 
may be one or two that are the females of this species, but not being 
as sure of their position as of these three males, they are left in the 
former group. 

Holotype in collection of Mr. E. L. Dickerson. 

Paratypes in collection of Doctor Ball and the Snow Collection. 
University of Kansas. 



PLATE VIII. 

Fig. 1. ..4. acuminatus. 

Fig. 2. A. brunneus. 

Figs. 3, 4, 5. A. inornatus. 

Fig. 6. A. productus. 

Fig. 7. A. viridis var. variegatus. 

Fig. 8. A. angulalus. 

Fig. 9. A. obtutits. 

Fig. 10. A. pallidus. 

Fig. 11. A. viridis. 

(130) 



Lawson: Genus Acinopterxs. 



131 



PLATE VIII. 




PLATE IX. 

Fig. 1. A. acuminatus. 

Fig. 2. A. productus. 

Fig. 3. A. angulatus. 

Fig. 4. A. pallidus. 

Fig. 5. A. hrunneus. 

Fig. 6. A. viridis. 

Fig. 7. ^. viridis var. variegatus. 

Figs. 8 and 9. A. inornatus. 



(132) 



Lawson: Genus Acinopterus. 



133 



PLATE IX. 








PLATE X. 

Fig. 1. A. acuminatus. 

Fig. 2. A. angulatus. 

Fig. 3. A. productus. 

Fig. 4. A. viridis. 

Fig. 5. A. ^midm. a^^J^iy<o 

Fig. 6. A. inornatus. 

V](i. 7. A. brunneus. 

(134) 



Lawson: Genus Acinopterus. 



135 



PLATE X. 






PLATE XI. 


Fig. 1. 


A. acuminatus 


Fig. 2. 


A. pallidus. 


Fig. 3. 


A. productiis. 


Fig. 4. 


A. angulatus. 


Fig. 5. 


A. viridis. 


Fig. 6. 


A. brunneus. 


Fig. 7. 


A. inornatus. 



(136) 






Lawson: Genus Acinopterls. 



137 



PLATE XI. 




PLATE XII. 

Fitis. 1 AND 2. A. acuminatus. 
Figs. 3 and 4. A. ohlulus. 
Figs. 5 and 6. A. angulatus. 
Figs. 7 and 8. A. viridis. 
Figs. 9 and 10. A. brunneus. 
Figs. 11 and 12. A. inornatus. 
Figs. 13 and 14. A. productus. 

(138) 



J 



Lawson: Genus Acinopterus. 



139 



PLATE XII. 

















THE 

KANSAS UNIVEKSITY 

Science Bulletin 



Vol. XIV, No. 5— October, 1922. 

(Whole Series, Vol. XXIV, No. 5.) 

ENTOMOLOGY NUMBER V. 



. CONTENTS: 

The Life History of the Toad Bug (Heteroptera), 

H. B. Hunger j or d. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post oflSce in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVEBSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 5. 



The Life History of the Toad Bug. 

Gelastocoris oculatus Fabr. (Gelastocoridse).* 

BY H. B. HUNGERFORD, 

Professor of Entomology, University of Kansas. 

INTRODUCTION. 

IN MY PAPER on "The Biology and Ecology of the Aquatic 
Hemiptera"t (pages 49-51) I gave the gist of what was known at 
the time concerning the habits and life history of the toad bug. Its 
habitat and feeding habits, together with a brief description of the 
ovum and fifth nymphal instar were given there. 

During the season of 1920 I had an opportunity to gather con- 
siderable data relative to these interesting insects. Mrs. Grace 
Wiley, a student of mine, knpwing of my desire to study Gelasto- 
coris, sent me a number of living adults from her home in Chanute, 
Kan., in the autumn of 1919. One of these I kept alive until Sep- 
tember, 1920. On May 14, 1920, she sent me a shipment of adults, 
and again on July 6 another small lot of the bugs. The live insects 
supplied by Mrs. Wiley thus made possible the notes here reported, 
and I wish to acknowledge my gratitude to her for her kindness. 

THE TECHNIQUE USED IN THE REARINGS. 

Tall stenders, or staining jars, of glass about the size of jelly 
glasses were used. In each of these was placed an inch of sand 
that had been sterilized by heat. The paired adults were confined 
in low stenders of various sizes, and the sand searched each day for 
eggs. The young were isolated in the tall stenders as soon as 
hatched, for they were cannibalistically inclined, and two young 

* Mr. Bueno recognizes this as a new species, G. acciduiis Ms. I confess I cannot dis- 
tinguish it from G. oculatus Fabr. 

t Kansas University Science Bulletin, vol. XI, Dec. 1919; 265 pages, 33 plates. 

(145) 



146 The University Science Bulletin. 

would get on but a few hours together. The sand was moistened 
each day, and the jars covered with ground-glass covers. Each 
nymph was examined daily for molts, which were removed and 
placed in vials of alcohol or on cotton in small tin boxes, and each 
instar skin of each insect reared was preserved separately for study. 
This has provided adequate material for study of structural details 
of each stage. 

We endeavored to determine whether the insects had any choice 
of soil, by placing them in pans containing sand on one side and 
sandy loam on the other. Our results were not conclusive. We 
also used sterilized sandy loam in some of the rearing jars instead 
of the sand and found it of no advantage. 

The insects were fed house flies, plant lice, oscinid flies, cicadellids 
and many other small insects taken in sweeping the grass upon the 
campus. Each day the dead carcasses were cleaned out of the rear- 
ing jars and freshly killed insects inserted. 

Mortality was very high, as a glance at the tables presented be- 
low will show, and indicates that some essential factor of their 
natural habitat was lacking. The fact that 116 nymphs out of 179 
died in the first stage, and that they usually succumbed on the 
date when molting might have been expected to occur, would point 
to a hazard of ecdysis. An examination of the dates of death of 
the older nymphs further substantiates this view. 

Toward the end of a stadium the nymph always appears plump 
instead of flat, and so the appearance of the nymph indicates ap- 
proximately its development. Thus some would become plump in 
a week and molt; others would develop more slowly. 

In spite of the discouragements because of the very high rate of 
mortality, the tending and study of the rearings were very interest- 
ing. Mr. William Hoffman, who assisted me very materially dur- 
ing the latter part of the summer, found the task most absorbing. 
He fed the nymphs and kept the records with as much interest, 
care and ingenuity as I could have done, and I desire to acknowledge 
herewith my indebtedness to him for his services. 

HABITAT. 

The toad bug is a shore bug, found along the muddy banks of 
small streams or the sandy beaches by the river. It is a notable 
example of protective coloration. Specimens taken on muddy banks 
are dull and slaty grey with indistinct pattern, while those from 
the sandy beaches are variegated, pebbled and mottled like the 
sand. Specimens in captivity have been observed to burrow be- 



Hungerford: The Toad Bug. 147 

neath tlie dirt in cloudy weather and remain tlius many liours. This 
habit may be their method of maintaining their geographical posi- 
tion in time of flood. I have observed broad, sandy, barren flats 
where toad bugs lived become inundated by rapid currents of 
water for a few hours; nevertheless, when the water receded and 
the sun came again, here were the bugs as before. There were 
neither vegetation nor sizable stones for their anchorage, so I sup- 
pose they "dug in." They hop about with considerable alacrity 
when alarmed (one first instar nymph jumped ten inches). They 
pounce upon their prey, which appears to consist of almost any 
sort of . insect they can capture, from a grouse locust (tettix) to a 
lacebug. 

MATING. 

There is considerable sexual dimorphism with the toad bug. The 
abdomen of the male is strongly asymmetrical, as is also the case in 
the male Corixidse. Figures illustrating this are shown on plates IX 
and XXXII of Science Bulletin, vol. XL At that time no ob- 
servations had been recorded on the mating. The male mounts the 
female, grasping her with the middle pair of legs, the first pair 
flexed beneath him, and the abdomen somewhat to the left side. 
This decided and invariable position to the left is due, of course, 
to the asymmetrical structure of the male genitalia. The fre- 
quency and duration of copulation of various pairs under obser- 
vation is given herewith. 



148 



The University Science Bulletin. 





MATING OBSERVATIONS COVERING 25 DAYS 












Pair 1. 


Pair 2. 


Pair 3. 


Pair 4. 


Pai 


r5. 


Pair 6 


Date. 


Obs. 
mat- 
ing. 


Hrs. 
dura- 
tion. 


Obs. 
mat- 
ing. 


Hrs. 
dura- 
tion. 


Obs. 
mat- 
ing. 


Hrs. 
dura- 
tion. 


Obs. 
mat- 
ing. 


Hrs. 
dura- 
tion. 


Obs. 
mat- 
ing. 


Hrs. 
dura- 
tion. 


Obs. 
mat- 
ing. 


Hrs. 
dura- 
tion. 


May 15 

May 17 


X 

X 

X 

X2 

X 

X 

X 

X2 

X 

X 

X 


4 + 
6+ 

2-1- 
2 

iV2+ 


X 

X 

X2 

X 

X 

X 


4-^ 

2 + 


X 
X 


4-f 


X 
X 
X 

X2 

X2 

X2 

X 

X 


4+ 
4+,2 


X 
X 
X 
X 


4+ 
6+ 
1 

l}/2 


X 


4+ 


May 18 






May 19 


2+ 

2H+ 

5-t- 


X 


2+ 


X 
X 

X 
X 

X2 


1-1- 


May 20 


6+ 


May 21 








X 


4H+ 


2 


May 22 








May 24 




X 


1 


X 
X 


V4 




X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 






May 25 




May 27 










X 






X 




May 28 




X 
X 




X 






May 29 






- 








May 31 


X 
X2 
X 
X 














1-h 

2+ 


X 
X 
X 




June 1 


3M 


X 

X 


7+ 
2+ 












June 2 


X 
X 


4+ 


X 


2+ 


1M+ 


June 3 


June 4 




X 
X 














June 7 


X 










X 










June 9 










X 





In the table X signifie.s observed mating, 
plus sign indicates that the bags were mating 



If followed by the figure 2 it 
when first or last observed, or 



indicates two separate matings. The 
both. 



The above table indicates that the mating clasp may last as long 
as seven hours, that matings are frequent, often twice a day, and 
that they occur almost daily over a considerable period of time. 
Matings were observed in the laboratory from May to November. 

OVIPOSITION. 

The eggs are either completely or partly buried in the sand, ceph- 
alic end uppermost. The partly buried dry egg is white and very 
difficult to discover in dry sand. When it is moistened it changes to 
an amber or ferruginous, and with the sand takes on a deeper colored 
and glistening appearance, which further adds to its resemblance to 
a sand grain. A photograph of the egg upon a background of sand 
is shown on plate XIII. The number of eggs deposited by a female 
during the season must be 200 or more, because 192 nymphs and eggs 
have been recorded for one female, and it is quite likely that I did 



Hungerfokd: Tiik Toad Bug. 149 

not count the full coniplenicnt. Oviposition continues throughout 
the season, the number of eggs laid per day varying from 1 to 13, 
from 2 to 6 being perhaps the average. A study of the hatching 
dates on the life-history tables will indicate the rate of egg-laying 
very well. 

INCUBATION. 

The incubation period varied from twelve to fifteen days. The 
red eyespots of the embryo are evident a number of days before 
hatching. The egg increases in size somewhat as the embryo devel- 
ops, and the egg becomes darker with development. 

HATCHING. 

I was fortunate enough to observe the hatching process with the 
binoculars on several occasions, but never in as satisfactory lighting 
as I should have liked. The cephalic end of the egg shell splits longi- 
tudinally, squarely between the eyes, and extends back above the 
dorsum of the embryo. Through this rent a white, bulging body 
appears, resembling the bubble found above the head of Corixa. 
The front part of the head of the embryo pulsates rapidly. By slow 
straining heaves the embryo crowds out through the opening. Its 
body is as soft and pliable as a caterpillar. By bulging the fore part 
of the body and contracting the latter part, it worms its w^ay to free- 
dom — a creamy-white creature marked wdth two large, dark-red 
eyes, and v/ith a body nearly cylindrical in shape and the thick 
limbs all most economically tucked away upon its venter. Then 
comes the postnatal molt, the casting of the shroud that binds the 
embryo. Standing erect upon its caudal end, its body encompassed 
and its limbs tied down by a diaphanous membrane that still holds 
it helpless to the empty casque in which it was formed, it struggles 
for freedom. First the membrane gives way above the head and 
emergence begins. As this skin slips back, the knob-like antennse, 
which were directed downward along the beak, are free and change 
their position. Then the beak appears, and after slowly bending 
back and forward, one front leg pops out free, then the other. Fi- 
nally all the legs are free, and the little bug settles down upon them, 
the shroud still about the tip of the abdomen. He flattens out into 
a toad bug, and after resting for a time, as if the birth struggles had 
been most exhausting, he suddenly becomes lively and starts away. 
Over the first moist pebble he passes, the molt is left, if by chance it 
did not remain fastened in the slit of the egg-shell. A period of 
thirty minutes often is consumed in the hatching. One bug that 



150 The University Science Bulletin. 

hatched at 2:35 p. m. was creamy white with dark-red eyes at 3:12 
p. m., when a faint pattern began to appear. At 3:35 it was some- 
what darker and the pattern more distinct. Thus it takes an hour 
or so for it to attain its characteristic color. 

MOLTING. 

An examination of the life-history tables will indicate that mor- 
tality at molting time was very great in the rearings. This suggests 
that conditions were not favorable to normal molting. In the light 
of R. Takahashi's observations on the Ochteridae, published in Jap- 
anese,* it may be that Gelastocoris nymphs, like those of Ochterus 
formosanus Mats., normally fashion for themselves small cells of 
sand above the ground in which the moltings take place. The sand 
in the rearing jars was packed, and perhaps too coarse for the 
nymphs. 

The nymph becomes very plump of body along toward the time 
to change. Several first-instar bugs were under observation during 
the process of molting. The bug rests upon the sand, all legs out- 
spread and apparently rigid. A longitudinal rent appears on the 
dorsum of the head and thorax, and the greenish or creamy-white 
nymph begins bulging out, the dorsal part of the thorax leading. 
Shortly the head is freed, the slit through the old skin extends back 
to the abdomen and then laterally to the margins of the body. These 
lateral fissures enable the bug to work his way out. The new form 
is so much larger than the old exuvium which encased it, that one 
wonders how it was ever tucked away in so small a fepace. When 
the new instar is entirely free, the old skin may snap back in place 
again and look like a perfect nymph, save that the eyes are whitish 
instead of dark red. 

The following article appeared in Japanese, by R. Takahashi. 
The observations are so interesting, and relate to a family so little 
known, that English-reading students will be glad to give Mr. Taka- 
hashi credit for it. 

These observations pertain to Ochterus jormosanus (Mats.), wliich is not 
uncommon in Formosa. 

1. The adults Hve upon the sandy shores of ponds and streams, where the 
colors of their backs merge into their surroundings, rendering them difficult to 
discover. They are not able to submerge and do not run out upon the water, 
where they are occasionally found by accident; but the nymphs are amphib- 
ious, being often seen submerged. 

* R. Takahashi: "Observations on the Ochteridte," Trans. Nat. Hist. Soc. Formosa, vol. 
XI, No. 55, pp. 119-125 (1921). 



Hungerford: The Toad Bug. 151 

2. The adults arc very active, while the nymphs are rather inactive. The 
death ft^igning has never been observed. 

3. The species is not gregarious, but two or three nymphs are sometimes 
found in groups. 

4. The n>-inphs sometimes \ibrate the abdomen vertically for a few seconds 
when resting on the shore. 

5. The nymphs cover their backs completely with sandy granules. All the 
nymphal instars have this habit. Their heads are provided on the front with 
12 to 14 stout processes jM'ojecting forward and arranged in a transverse row, 
with which they scoop the sand upon the heads and push them backward by 
the front legs. 

6. The m-mphs construct for themselves small cells of sand above the 
groimd, using the processes on the front, in which the meltings take place. 
All the instars have this peculiar habit. 

7. The nymphs with the dorsums wettable are amphibious in habit, being 
often found submerged. When submerged, the bodies are held always just 
below the surface film, and they swim rather awkwardly, moving all the legs, 
but do not swim deeper. A store of air for respiration when submerged, is 
carried with the insect on the lower surface of the abdomen, and the nymphs 
now and then turn on their backs at the surface, thus exposing the lower sur- 
face of the abdomen into free air to take new supply of air. This is done very 
quickly. 

8. The mating habit is almost as in the insects of Microvelia, but the 
males do not remain on the backs of their mates for a long time when the 
copulation is finished. The males and females mate repeatedly. 

9. The eggs are singly placed upon the sandy granules, or upon the decayed 
leaves on the ground. 

10. The egg is similar in shape to that of Gelastocoris, species figured by 
Doctor Hungerford (1919), measuring about 0.7 mm. in length. 

11. There are five nymphal instars, as is common for other Heteroptera, 
and the nymphal stage lasts more than one month. 

12. In the adults the front and middle tarsi are two-jointed, and the hind 
three-jointed, while in the nymphs all the tarsi are always two-jointed. 

13. The adults may be seen near Taihoku at almost any time. 

NOTES ON REARINGS. 
FIRST PAIR. 
Rearing Number 4- 
This pair consisted of the female that I had kept in the laboratory- 
all winter and a male selected from the spring shipment. They 
were confined in a six-inch stender in which had been placed a layer 
of sand with a place scooped out on one side for water. The water, 
however was taken up by the sand and the entire mass became 
water-soaked. To make a dry footing for tlie bugs, a bit of cork 
was set upright in the soil, but this too became wet. Green algae 
covered the sand, the cork, the sides of the jar, and even the bugs 
were green with it. The surface of the sand was found on July 1 



152 The University Science Bulletin. 

to be teeming with nematodes tliat were intent upon consuming 
the fly carcasses supplied as food for the bugs: No careful examina- 
tion was made of this jar until July 1, when four white eggs were 
found on top of the cork, placed there in an endeavor, apparently, 
to keep them out of the wet, soggy sand. I then supplied dry sand 
to reduce the moisture, and on July 6 found eggs in tlie sand; 
observed mating on that date also. July 12 two nymphs hatched. 
July 20, there were several active first-instar nymphs. July 24, 
removed 18 dead first-instar forms and observed nine live ones. 
There were several eggs yet to hatch and some fresh eggs. July 27, 
removed two second-instar forms and took out 18 dead first-instar 
bugs. Seven live ones were observed at this time. July 30, observed 
a couple hatch and counted 10 active first-instar nymphs; also re- 
moved 7 dead ones. On August 1 there were nine live first-stage 
forms; removed 2 dead ones. August 2, counted 11 live first-stage 
bugs and removed 6 dead ones. On the 3d removed 2 dead ones. On 
the 4th there were 3 newly hatched bugs, and I removed 5 dead 
ones. On August 5 one first-instar bug was observed feeding upon 
another. August 7 there were at least 18 nymphs, three of them 
white, denoting recent hatching. August 8, 7 dead firsts were re- 
moved, but a dozen were still lively. August 9 another freshly 
emerged nymph was noted, and on the 10th 2 more. August 13, 
removed a second-instar form, and on August 15 another newly 
hatched was noted. August 16, a dark nymph was caught in the 
act of killing a white newly emerged brother. He was upon the 
back of his victim with beak inserted just back of the unfortunate's 
head. The latter was struggling, but to no avail. On August 18 
one second-instar form w'as isolated and 34 dead first-stage bugs 
removed. Mr. Hoffman found three nymphs feeding upon their 
fellows, and all three victims were alive and kicking. There were 
about a dozen live bugs of this stage. August 21' brought forth 
3 second-stage bugs, which were taken out. August 22, 11 dead 
firsts were moved, and on the 23d 3 dead firsts and 1 live second 
removed. August 24, 2 dead firsts were taken out. August 29, 5 
dead first were removed, and on August 31 the female was found 
dead. Twenty-three dead nymphs were counted out and 8 eggs 
transferred to another stender. No live nymphs were present, and 
the jar was set aside till September 20, when a careful count was 
made of remains — 10 eggs and 16 nymphs completed the record for 
this pair. The above notes have been given to show the result of 
trying to rear the bugs together. 



Hungerford: The Toad Bug. 



153 



This pair of bii^s was observed mating May 15, 17, 18, 19, 20, 21, 
22, 24, 27, June 2, 7, 22, and July 6. No count of eggs laid was 
attempted, but 22 eggs, 160 first-instar and 10 second-instar nymphs 
were taken, a total progeny of 192, with egg-lay noted from July 1 
to last of August — this the performance of a female that spent the 
preceding winter in the laboratory. 









REARING 


RECORD OF NO. 


4. 












No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


1 

Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


4a 


July 12 


July 21 


Aug. 2 


Aug. 15 


Aug. 30 


Sept. 19 


& 


9 


12 


13 


15 


20 


69 


Oct. 6 


4b 


July 12 


July 20 


Died 
July 24 










8 


4° 
































4c 


July 16 


July 24 


Aug. 13 


Died 
Aug. 16 








8 


20 


3° 




















.... 






4d 


July 16 


July 25 


Aug. 5 


Aug. 16 


Aug. 31 


Sept. 23 


cf 


9 


11 


11 


15 


24 


70 


Nov. 17 


4e 


Aug. 1 


Aug. 13 


Died 
Aug. 31 










12 


18° 
































4f 


Aug. 8 


Aug. 18 


Died 
Aug. 31 










10 


13° 
































4g 




Aug. 20 


Sept. 10 


Died 
Sept. 14 










21 


4° 




























4h 




Aug. 21 


Sept 11 


Sept. 20 


Oct. 5 


Died 
Oct. 20 






21 


9 


16 


15° 


71? 












4i 




Aug. 21 


Sept. 6 


Sept. 17 


Sept. 27 


Oct. 26 


& 




16 
1° 


11 


10 


29 


76? 


Oct. 28 


4j 




Aug. 23 


Died 
Sept. 24 



















° indicates "died" in this and following tables. 

SECOND PAIR— OFFSPRING IN FOUR SERIES. 
Rearing Number 1, Series m. 
On July 6, Mrs. Wiley brought me 7 bugs from Chanute, which 
she had captured July 3. One pair, observed mating, I placed in a 
small stender on sand with a sprig of moss. July 8, quite a number 
of eggs were in the sand, so the adults were removed to another 
stender. On July 15, the eggs showed pink eyespots of the embryo 
within; and on July 19, 8 hatched and were isolated for rearing. 
July 21, 5 more hatched; and July 24, 3 more. Thus at least 16 
eggs were laid by this female in two days, and the incubation period 
was 12 or 13 days. The rearing table marked Im series is a record 
of fifteen of these nymphs. 



154 The University Science Bulletin. 

Rearing Number 1, Series n. 
The pair was transferred from Im to this stender, containing 
sand, on July 8 ; observed mating on the 11th and removed on the 
12th. Could find no eggs, and though this stender was studied till 
July 29, no nymphs appeared. 

Rearing Number 1, Series o. 
Transferred the pair from number In to this stender, containing 
sand, July 12. On July 24, 4 nymphs hatched, and so I removed the 
adults to another stender and transferred the nymphs to jars as they 
hatched. The rearing table marked lo series indicates the dates of 
hatching. But to this list of 49 must be added the following: July 
27, 6 first instars in large stender, labeled lOh. August 6, 4 dead 
instars; and August 8, 2 more. A total, therefore, of 61 hatched 
from eggs deposited between July 12 and 24, an average of 5 per 
day, with a record of 13 for August 6. Incubation period, 12 days. 

Pair Number Ip. 

This pair of adults was transferred from lo on July 24. They 
were observed mating on this date and again on August 8, when 2 
first-instar forms hatched. Removed the male on August 10 and 
returned on the 18th, This female was caught twice feeding upon 
her own offspring. Besides the 65 nymphs hatched from August 8 
to September 27 and used in rearings, 28 dead ones were taken out 
of this stender in which the pair was confined — a total of 93 off- 
spring between July 24 and September 27. The observed matings 
were on July 24, August 8, August 21 and September 20. One adult 
died October 4 and the other October 23. The rearing data are pre- 
sented on that portion of the table marked IP series. 

Summing up for this pair of bugs taken from the wild on July 3 
and entered for observation on July 6, we get a total progeny of 170 
hatched, and of this number we were able to rear to the adult stage 
4 insects. 



Hungerford: The Toad Bug. 



155 



SECOND PAIR, REARING TABLE. 

Series Im. 



No. 


Date 

hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


Inift 


July 19 


Died 
Aug. 14 












26° 






































Imb 


July 19 


Died 
Aug. 7 












19° 








































July 19 


Aug. 11 


Oct. 7 


Died 
Oct. 25 








23 


27 


19° 


























Imd 


July 19 


Aug. 6 


Died 
Aug. 12 










18 


6° 
































Ime 


July 19 


Died 
Aug. 5 












17° 






































Imf 


July 19 


Died 
Aug. 9 












21° 






































Img 


July 19 


Died 
July 28 












9° 




































Imh 


July 21 


Aug. 1 


Aug. 11 


Died 
Aug. 16 








11 


10 


5° 


























Ipii 


July 21 


July 28 


Aug. 11 


Died 

Aug. 16 








7 


14 


5° 


























Imi 


July 21 


Aug. 7 


Died 
Aug. 11 










17 


4° 
































Imk 


July 21 


Died 
Sept. 12 












53° 






































1ml 


July 21 


Died 
July 30 












9° 






































Imm 


July 21 


July 25 


Aug. 6 


Aug. 15 


Aug. 30 


Sept. 19 


cf 


4 


12 


9 


15 


20 


60 


Nov. 4 


Imn 


July 24 


Aug. 9 


Died 
Aug. 10 










16 


1° 
































Imo 


July 24 


Died 
July 30 












6° 
















i 





156 



The University Science Bulletin. 



Series lo. 



No. 


Date 

hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


loa 


July 24 


Aug. 11 


Died 

Aug. 14 










18 


2.° 
































lob 


July 24 


Aug. 10 


Died 
Aug. 13 










17 


3° 


































July 24 


Died 
July 30 












6° 






































lod 


July 24 


Died 
July 31 












7° 








































July 25 


Died 
Aug. 10 












16° 






































lof 


July 25 


.Aug. 12 


Sept. 2 


Sept. 17 


Died 
Oct. 1 






18 


21 


15 


13° 




















log 


July 25 


Aug. 10 


Died 

Aug. 14 










16 


4° 






























loi 


July 29 


Died 

Aug. 8 












10° 






































loj 


July 29 


Died 
Aug. 5 








- 




7° 




































lok 


July 29 


Aug. 13 


Aug. 31 


Sept. 13 


Sept. 25 


Oct. 21 


9 


15 


18 


13 


12 


26 


84 


Dec. 20 


lol 


July 29 


escaped 




















































lom 


July 29 


Died 
.Aug. 11 












13° 








































July 29 


Died 

Aug. 5 












7° 


































July 29 


Died 
Aug. 10 












12° 






































lop 


July 30 


Died 
Aug. 7 












8° 




































loq 


July 30 


Aug. 13 


Sept. 15 


Died 

Sept. 24 








14 


33 


9° 
























lor 


July 30 


Died 
Aug. 1 












2° 






































los 


July 30 


Aug. 11 


Aug. 28 


Sept. 12 


Sept. 24 


Died 
Oct. 22 




12 


17 


15 


12 


29 


85 




lot 


July 30 


Died 

Aug. 28 












29° 








































July 30 


Died 

Aug. 8 












9° 








































July 30 


Died 

Aug. 8 












9° 






































lox 


July 30 


Died 

Aug. 9 












10° 






































loy 


July 30 


Died 

Aug. 8 












9° 




































loaa 


Aug. 1 


Died 
Aug. 9 












8° 















Hungerford: The Toad Bug. 



157 



Series lo. — Concluded. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. - 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


lobb 


Aug. 1 


Aug. 18 


Died 
Sept. 17 










17 


30° 
































locc 


Aug. 1 


Died 
Aug. 8 












7° 






































lodd 


Aug. 2 


Died 
.\ug. 10 












8° 








































Aug. 2 


Died 
Aug. 9 












7° 






































loff 


Aug. 2 


Died 
Aug. 9 












7° 






































logg 


Aug. 2 


Died 
Aug. 9 












7° 




































lohh 


Aug. 2 


Died 

Aug. 9 












7° 




































loii 


Aug. 2 


Died 
Aug. 9 












7° 






































lojj 


Aug. 2 


Died 
Aug. 9 












7° 




































lokk 


Aug. 4 


Died 
Aug. 10 












6° 






































loll 


Aug. 4 


Died 
Aug. 14 












10° 








































Aug. 4 


Died 
Aug. 9 












5° 








































Aug. 4 


Died 
Aug. 9 












5° 








































Aug. 5 


Died 
Aug. 10 












5° 






































lopp 


Aug. 5 


Died 
Aug. 15 












10° 




































loqq 


Aug. 5 


Sept. 12 


Sept. 26 


Oct. 21 


Died 
Nov. 17 






38 


14 


25 


27° 




















Aug. 6 


Died 
Aug. 11 












5° 








































Aug. 6 


Died 
Aug. 10 












4° 






































lott 


Aug. 6 


Died 
Aug. 13 










7° 








































Aug. 6 


Died 
Aug. 9 












3° 




































Aug. 6 


Died 
Aug. 12 












6° 








































Aug. 6 


Died 
Aug. 1-1 












8' 








































Aug. 6 


Died 
Aug. IC 












4" 
















. 
























Aug. 6 


Died 
Aug. IC 












7' 
















. 






















lobbb 


Aug. 6 


Died 
Aug. 1 


. 










5' 













158 



The University Science Bulletin, 



Series Ip. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


Ipa 


Aug. 8 


Died 
Aug. 14 












6° 






































Ipb 


Aug. 8 


Died 
Aug. 14 












6° 






















^ 
















Idc 


Aug. 9 


Died 
Aug. 14 












5° 






































Ipd 


Aug. 9 


Died 
Aug. 15 












6° 








































Aug. 9 


Sept. 22 


Died 
Oct. 15 










44 


23° 
































lof 


Aug. 10 


Died 
Aug. 18 












8° 






































iDff 


Aug. 10 


Died 
Aug. 16 












6° 






































Iph 


Aug. 10 


Died 
Aug. 19 












9° 






































Ipi 


Aug. 10 


Died 
Aug. 19 












9° 




































Ipj 


Aug. 10 


Died 
Aug. 17 












7° 




































Ipk 


Aug. 11 


Died 
Aug. 12 












1° 




































Ipl 


Aug. 12 


Died 
Aug. 25 












13° 




































1pm 


Aug. 16 


Died 
Aug. 22 












6° 




































Ipn 


Aug. 16 


Died 

Aug. 22 












6° 

8° 




































Ipo 


Aug. 16 


Died 
Aug. 24 


















• 
























Ipp 


Aug. 16 


Died 
Sept. 9 












24° 




































Ipq 


Aug. 16 


Died 
Aug. 23 












7° 




































Ipr 


Aug. 16 


Died 
Sept. 2 












17° 




































Ips 


Aug. 17 


Died 
Aug. 26 












9° 




































Ipt 


Aug. 17 


Died 
Aug. 29 












12° 




































Ipv 


Aug. 18 


Sept. 13 


Died 
Oct. 6 










26 


23° 






























Ipw 


Aug. 19 


Died 

Aug. 1 












13° 




































Ipx 


Aug. 21 


Died 

Aug. 2 












12° 












• 
























ipy 


Aug. 21 


Sept. 9 


Sept. 23 


Oct. 11 


Died 
Oct. 23 






19 


14 


18 


12° 









Hungerford: The Toad Bio. 



159 



Series Ip — Continued. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


Ipz 


Aug. 22 


Died 
Aug. 11 












20° 




































Ipaa 


Aug. 23 


Died 

Aug. 4 












12 = 




































Ipbb 


Aug. 25 


Died 
.\ug. 16 












22° 




































Ipcc 


Aug. 26 


Died 
Aug. 29 












3° 




































Ipdd 


Aug. 27 


Died 
Sept. 4 












8° 




































Ipee 


Aug. 28 


Sept. 13 


Died 
Sept. 24 










16 


11° 
































Ipff 


Aug. 29 


Sept. 15 


Sept. 25 


Oct. 14 


Nov. 17 


Died 

Nov. 26 




17 


10 


20 


34 


9° 












Iphh 


Aug. 30 


Died 
Sept. 9 












10° 






































loii 


Sept. 3 


Died 
Sept. 17 












14° 


































Ipjj 


Sept. 4 


Died 

Sept. 24 












20° 






































Ipkk 


Sept. 4 


Died 
Sept. 15 












11° 






































IpU 


Sept. 7 


Died 
Sept. 9 












2° 







































Innun 


Sept. 7 


Died 
Sept 15 












8° 








. 




















Ipnn 


Sept. 9 


Sept. 21 


Oct. 15 


Nov. 4 


Died 
Dec. 4 






12 


24 


20 


30° 





















Ipoo 


Sept. 9 


Died 

Sept. 18 












9° 






































Ippp 


Sept 9 


Died 
Sept. 17 












8° 






































Ipqq 


Sept. 9 


Died 
Sept. 19 












10° 






































Iprr 


Sept. 10 


Died 
Sept. 21 












11° 






































Ipss 


Sept. 12 


Died 

Sept. 22 












10° 






































Iptt 


Sept. 12 


Died 
Sept. 26 












14° 






































Ipuu 


Sept. 12 


Died 
Sept. 21 












9° 






































Ipw 


Sept. 12 


Died 
Sept. 28 












16° 






































Ipww 


Sept. 12 


Oct. 8 


Died 
Oct. 12 










26 


4° 
































Ipxx 


Sept. 12 


Sept. 23 


Oct. 11 


Died 

Nov 13 








11 


18 


33° 











160 



The University Science Bulletin. 



Series Ip — Concluded. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 

molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days in stages. 


Adult 


1st. 
10° 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


Ipvy 


Sept. 13 


Died 
Sept. 23 




































Ipzz 


Sept. 13 


Died 
Sept. 21 












8° 






































Ipaaa 


Sept. 13 


Sept. 29 


Died 
Nov. 2 










16 


3° 
































Ipbbb 


Sept. 13 


Died 
Sept. 28 












15° 






































Ipcee 


Sept. 15 


Oct. 5 


Nov. 2 


Died 
Dec. 30 








20 


28 


59° 


























Ipddd 


Sept. 15 


Died 

Sept. 25 












10° 






- 
































Ipeee 


Sept. 15 


Sept. 27 


Died 
Oct. 15 










12 


18° 


































IpiTf 


Sept. 17 


Died 
Sept. 30 












13° 






































lD2Ee 


Sept. 17 


Died 

Sept. 25 












8° 












■• 


























Iphhh 


Sept. 17 


Died 
Sept. 19 












2° 






































Ipiii 


S?pt. 22 


Died 
Oct. 2 












10° 






































Ipjji 


Sept. 22 


Died 

Sept. 2S 












6° 








































Ipkkk 


Sept. 23 


Died 
? 












? 




































IpUl 


Sept. 23 


Died 
Oct. 13 












20° 






































Ipmmm 


Sept. 23 


Oct. 18 


Died 
Oct. 30 










25 


12° 
































Ipnim 


Sept. 25 


Dir-d 
Sept. 29 












4° 
4° 




































Ipooo 


Sept. 27 


Died 
Oct. 1 





























It was not planned when I began the studies to carry the rearings 
to such an extent and therefore the simple system of designating 
became cumbersome. 



Hungerford: The Toad Bug. 



161 



THIRD PAIR. 



On Alay 14 I placed a pair in a small stendcr on the sand. Eggs 
showed eyespots June 2. June 10, 4 nymphs appeared, and on this 
date adult female died. The rearings from this pair are given in the 
following table: 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days i n stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


lb 


June 10 


June 28 


Died 
July 17 










18 


19° 
































Ic 


June 10 


June 19 


July 1 


Escaped 
July 6 








9 
15° 


12 




















■- 










Id 


June 13 


Died 
June 28 










































le 


June 13 


Died 
June 21 












8° 






































If 


June 14 


June 24 


July 3 


July 11 


July 19 


Aug. 2 


9 


10 


9 


8 


8 


12 


47 


Jan. 11, 
1921 


Ih 


June 17 


June 28 


Died 
June 28 










11° 




































li 


June 17 


July 1 


Died 
July 6 










14 


5° 
































Ij 


June 21 


Died 

July 2 












11° 






































Ik 


June 21 


June 28 


July 6 


July 15 


July 22 


Aug. 7 


9 


7 


8 


9 


7 


16 


47 


Nov. 2 


11 


June 22 


July 5 


July 15 


July 23 


Aug. 6 


Aug. 31 


_9_ 


13 


10 


8 


14 


25 


70 


Oct. 10 



162 



The University Science Bulletin. 



FOURTH PAIR. 

In a large stender containing sandy loam, and labeled No. 7, was 
placed a mating pair of bugs, June 28. Eggs were discovered in the 
soil on July 8. July 13 nymphs hatched and the adults were re- 
moved. July 20, 7 more nymphs were out. These 12 nymphs were 
isolated in stenders and a tabular report of these follows. The in- 
cubation period was somewhere between 7 and 15 days. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


7a 


July 13 


Escaped 




















































7b 


July 13 


July 20 


Died 
July 23 










7 


3° 
16 






















8° 










7c 


July 14 


July 21 


Aug. 6 


Died 

Aug. 14 








7 
9° 


























7e 


July 14 


Died 
July 23 










































7f 


July 20 


Died 
July 30 












10° 






































7g 


July 20 


July 28 


Aug. 6 


Aug. 16 


Sept. 1 


Sept. 21 


9 


8 


9 


10 


16 


20 


63 


Oct. 28 


7h 


July 20 


Aug. 16 


Aug. 30 


Sept. 12 


Sept. 22 


Oct. 17 


cf 


27 


14 


13 


10 


25 


89 


Nov. 14 


7i 


July 20 


Aug. 4 


Aug. 14 


Aug. 26 


Sept. 11 


Sept. 29 


& 


15 


10 


12 


16 


18 


71 


Nov. 21 


7j 


July 20 


Aug. 9 


Died 
Aug. 12 










20 


3° 
































7k 


July 20 


Died 
July 28 












8° 




































71 


July 31 


Aug. 6 


Aug. 16 


Sept. 14 


Died 
Sept. 27 






6 


n 


29 


13° 

















Hungerford: The Toad Bug. 



1H:} 



FIFTH SERIES. 

A sprig of moss and eartli containing eggs were plaeed in a slender 
July 12. Tiiese eggs were deposited by bugs sent rae July 6, amongst 
the material in which they were packed. They began hatching July 
20 and finished July 24. The record of the eight nymphs isolated is 

sliown in tabular form below. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 




Days in stages. 


Adult 


1st. 


2i 


3d. 


4th. 


5th. 


Tot. 


died. 


9a 


July 20 


July .30 


Aug. 19 


Sept. IP 


Oct. 7 


Died 
Oct. 18 




10 


20 


27 


22 


11 












9b 


July 20 


Aug. 6 


Aug. 18 


Aug. 30 


Sept. 13 


Oct. 5 


& 


17 


12 


12 


14 


22 


77 


Oct. 17 


9c 


July 21 


Aug. 2 


Died 
Aug. 10 










12 


8 
































9d 


July 21 


Died 
July 31 












10 






































9e 


July 24 


Died 
Aug. 9 












16 






































9f 


July 24 


Died 
Aug. 3 












10 






































9g 


July 24 


Aug. 1 


Died 
Aug. 9 










8 


8 






















.... 










9h 


July 24 


July 30 


Aug. 15 


Sept. S 


Sept. 22 


Died 
Oct. 14 




6 


16 


25 


13 


22 







164 



The University Science Bulletin. 



SIXTH PAIR. 

Placed 2 female bugs in large stender marked No. 12, on July 20. 
These bugs were from those Mrs. Wiley collected July 3. On 
August 2, 2 nymphs appeared — an incubation period of 13 days. 
The nymphs continued to appear, and their record is shown in 
table 12 below. September 5 only 1 adult was to be found, and 
yet it was impossible for the bugs to escape from the jar. Septem- 
ber 6 both were gone from view, but later they were again above 
the sand. In cloudy, threatening weather, the toad bugs bury 
themselves beneath the surface and appear again in sunny weather. 

Table 12. 



No. 


Date 
hatched. 


1st 
molt. 


2d 
molt. 


3d 
molt. 


4th 
molt. 


5th 
molt. 


Sex. 


Days in stages. 


Adult 


1st. 


2d. 


3d. 


4th. 


5th. 


Tot. 


died. 


12a 


Aug. 2 


Died 
Aug. 8 












6 






































12b 


Aug. 2 


Died 
Aug. 9 












7 






































12c 


Aug. 8 


Died 

Aug. 14 












6 






































12d 


Aug. 9 


Died 

Aug. 14 












5 




































12e 


Aug. 10 


Died 
Aug. 12 












2 






































12f 


Aug. 10 


Aug. 26 


Died 
Sept. 17 










16 


22 
































12g 


Aug. 11 


Died 
Aug. 19 












8 




































12h 


Aug. 15 


Died 

Aug. 21 












6 






































12i 


Aug. 15 


Died 

\ug ?2 












7 








































12j 


Aug. 16 


Died 

Aug. 28 












12 









SEVENTH PAIR. 

In stender marked 2a, placed a mating pair on May 25. On June 
3, found an egg imbedded in the sand for about one-half of its 
length. On June 13, found 11 eggs, 5 or 6 of which showed pink eye- 
spots, so removed the adults to another stender. The eggs began 
hatching June 17 and continued until June 28. The minimum in- 
cubation period for some of the eggs was 15 days. 



Hungerkord: The Toad Bug. 165 

EIGHTH PAIR. 

Pair from Chanute placed in stcnder labeled la, June 2. The fe- 
male died June 10. Eggs hatched June 24 and June 28. Incuba- 
tion period at least 14 days, likely longer. 

NINTH PAIR. 

Female emerged from one of the rearings (11) on August 31, from 
a nymph that hatched June 22. Added a male, but no mating ob- 
ser\ed, and female died October 10. 

TENTH PAIR. 

Female emerged from one of the rearings (Ik) on August 10, from 
nymph that hatched June 21. Placed a male with her and matings 
were observed on August 13, 15 and 18. No eggs w^ere found after 
most thorough search. Female died November 4. 

DESCRIPTION OF STAGES. 

(See plates XIII and XIV.) 

THE EGG. 

Size. Length, 1.25 mm.; diameter, .91 mm. 

Shape. Broadh' oval. The sui-face is roughly granular and marked into 
regular hexagonal areas bj' thickened ridges of the chorion. 

FIRST-INSTAR NYMPH. 

Size. Length, 2 mm.; width of thorax, 1.4 mm.; width of head, 1.2 mm. 

Color. Eyes dark; body mottled to checkered obscurely, general color 
sometimes light, sometimes dark. Legs and abdomen are usually marked a^ 
described for later instars. 

Structural peculiarities. Eyes larger relatively than in succeeding instars. 
They are not placed upon as high elevations and the inner emarginations are 
not marked. The beak is four-segmented; the antenuEe three-segmented; and 
the tarsi appear one-segmented, terminating in two claws. The middle and 
hind tarsi, however, have a short basal segment, making two. Lateral margin 
of mesothorax less in length than that of metathorax. 

SECOND-INSTAR NYMPH. 

Size. Length. 2.5 mm.; width of thorax, 1.9 mm.; width of head, 1.5 mm. 

Color. Eyes still dark, not distinctly banded. Pattern obscured, that of 
limbs and abdomen as in later instars. 

Structural peculiarities. The inner emargination of the eyes a little more 
marked than in the preceding instar. Lateral margin of mesothorax equal in 
length to that of metathorax. 



166 The University Science Bulletin. 

THIRD-INSTAR NYMPH. 

Size. Length, 3.5 mm.; width of thorax, 2.7 mm.; width of head, 2.0 mm. 

Color. The eyes faintly banded with three bars of color, one of them on 
the inner margins. The general pattern variable; but when defined, like 
other instars. 

Structural ])eculiarities. The inner emargination of the eye now regular 
and marked. Ocelli faintly visible in some, while in others unmistakably 
present. Lateral margin of Mesothorax now a little longer than that of 
metathorax. 

FOURTH INSTAR NYMPH. 

Size. Length, 4.5 mm.; width of thorax. 3i.6 mm., width of head, 2.4 mm. 

Color. The general pattern as in fifth. The eyes banded by four faint bars, 
one of which is on inner margin. 

Sttructural 'peculiarities. The lateral margin of mesothorax now is about 
twice as long as that of the metathoracic margin, due, of course, to the 
lengthening of the wing pads. The body is covered with closely set short, 
stout, appressed spines, which show more plainly than in the preceding instar. 
The ocelli show plainly now. They are located just above the lateral arms 
of what I take to be the epicranial suture. The nymphal exuvium always 
shows a Y-shaped rent on the head, and it is on the ui^per margin of the 
lateral arms of this fissure that the ocelli are located. 

FIFTH INSTAR NYMPH. 

Size. Length, 6.2 mm.; width of thorax, 5.0 mm.; width of head, 3.0 mm. 

Color. Color variable from very light to very dark, and the pattern varying 
from obscure to distinct. The eyes have five dark bars. There are two pairs 
of black dots on the face, one above the other, and laterad and below the 
ocelli. A median black dot on vertex. The pronotum has front two-thirds 
of lateral margins of prothorax darker. There is a pair of black spots caudo- 
laterally of each pronotal elevation, of which there are two. The mesonotum 
has two rectangular black spots on front margin either side of median line. 
There is a transverse row of four black dots across caudal third of mesonotum; 
wing pad has a dark rectangular spot on shoulder; the middle of the pad 
crossed by another dark area; tip and inner margin of pad dark. Two faint 
dark irregular spots in the outer third of wing pad gives a mottled effect to 
the whole. A white spot is found on second abdominal segment either side 
of median line. A row of dark marginal spots, roughly triangular, cover the 
front two-thirds of the margin of each abdominal segment. There are also 
two rows of dark submarginal spots, faint and ill defined. The legs are banded 
with dark bands. The hind tibise have three dark bands besides being dark 
at the ends. 

Structural -peculiarities. The wing pads of the mesothorax now extend 
almost to caudal margin of thorax, nearly obscuring lateral margin of meta- 
thorax. 



Hungerford: The Toad Bug. 167 



SUMMARY. 

The toad bug places its eggs in the sand. The eggs hatch in about 
12 days. There are five nymphal instars. Forty-nine first-instar 
bugs transformed in an average of 15 days (the shortest time 4 
days and the longest 44 days). Thirty-three second-instar bugs 
transformed in an average of 16 days (shortest time 8 days and long- 
est 33 days). Twenty-two third-instar bugs averaged 15 days (the 
shortest time 8 days and the longest period, not counted in the 
average because it failed to molt, was 59 days). Eighteen fourth- 
instar forms averaged 15^/^ days (minimum 7 days and maximum 
34 days). Thirteen fifth-instar forms averaged 22 days (minimum 
12 and maximum 29 days). The average number of days for the 
thirteen adults to develop from the hatching to emergence was 701/2 
days. The shortest period was 47 days and the longest 89 days. 
By adding the 12 days incubation of the egg, we get a total de- 
velopment period of from 60 to 100 days. The adult female may 
deposit a dozen eggs a day, but would average perhaps only 2 to 6 
over the period of two or more months. One hundred and ninety- 
two eggs and nymphs were counted from one female from July 6 
to September 27. The nymphs possess ocelli, plainly distinguished 
as early as the third instar. All the stages are predaceous and much 
like their parents in other habits as well. 



168 The University Science Bulletin. 



PLATE XIII. 

Gelastocoru oculatus Fabr. 

Fig. 1. Adult bug (not oculatus, but an undescribed species in western 
Kansas) upon sand, showing how its mottled pattern makes it difficult to 
discern. 

Fig. 2. Ventral view of above species of bug. Enlarged photograph show- 
ing the bug clasping three lace bugs. 

Fig. 3. First-instar nymph, Gelastocoris oculatus Fabr. Enlarged dorsal 
view, same species showTi in figures 1 and 2. 

Fig. 4. Another toad bug. Enlarged ventral view, showing bug with prey. 

Fig. 5. First-instar nymi)h, Gelastocorifi oculat^is Fabr., ventral view. 

Fig. 6" Microphotograj)!! of egg and first-instar nym])h. Nymph in center 
of picture and the white oval egg to the right of it. The egg resembles the 
sand grains very closely. 



PLATK XIII. 






(169) 



PLATE XIV. 

Gelastocoris ociilatii.'< Fabr. 

Fig. 1. Egg. 

Fig. 2. First-instar nymph. 

Fig. 3. Second-instar nymph. 

Fig. 4. Third-instar nymph. 

Fig. 5. Fourtli-instar nymph. 

Fig. 6. Fifth-instar nymph. 

(170) 



.?■• 



Hungerford: The Toad Bi(;. 



171 



PLATE XIV. 




THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 6— October, 1922. 

(Whole Series, Vol. XXIV, No. 6.) 

ENTOMOLOGY NUMBER V 



CONTENTS: 

A New Subterranean Isopod (Crustacea)...//. B. Hungerford. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the po.st office in Lawrence as second-cla.ss matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

^■oL. XIV.J October, 1922. [No. 6. 



A New Subterranean Isopod from Kansas. 

Ccecidotea tridentata (Crustacea). 

By H. B. HUNGERFORD, 

Professor of Entomology, University of Kansas. 

IN MARCH of 1919, Mr. William Hoffmann, field assistant in our 
department of entomology at the University of Kansas, brought 
to me for determination some specimens of an isopod which he 
had taken from a cistern in Lawrence, Kan. 

They prove to belong to a new species of the genus Coecidotea. 
For them I propose the name Coecidotea tridentata, because the 
propodus of the first pair of legs of the male is armed with three 
conspicuous processes, a character which separates them from the 
previously described species. 

The crustacean genus Coecidotea Pack., as the name implies, is 
characterized by the absence of eyes, by the fact that the terminal 
segment of the body is much longer than broad, and by the elongate, 
narrow body. An analytical key to the genus was given by Harriet 
Richardson in her monograph of the isopods of North America, in 
1905. There were known at that time four species, namely, C. stygia 
Pack., C. nickajackensis Pack., C. richardsonce Hay and C. smithsii 
Ulrich. Doctor Ortmann, 1918, in chapter XXV of "Fresh-water 
Biology," had Miss Richardson's work in mind when he said there 
were four species of the genus and that they are found in caves, 
springs issuing from caves, and artesian wells. However, in 1911, 
in the Pomona College Journal of Entomology, volume 3, No. 3, 
Blanche Stafford described a fifth species, namely, Coecidotea ala- 
bamensis, from a well in Auburn, Ala. 

The following table will serve to separate the six species of the 
genus now known: 

(175) 



176 The University Science Btlletin. 

A. Propodus of first pair of legs armed witli one or more triangular processes. 

B. Propodus of first pair of legs armed with a triangular process near 
the distal end and with a long spine at the proximal extremity. 
Uropods about one-half the length of terminal abdominal segment. 
Outer branch three-fourths as long as inner, which equals the 
peduncle in length. C. nickajackennis Pack. 

BB. Propodus of first pair of legs armed with two triangular processes. 

C. Propodus with three additional short processes. Uropods 
about as long as terminal abdominal segment. Outer branch 
two-thirds as long as inner, which is two-thirds as long as 
jieduncle. C. stygia Pack. 

CC. Propodus with three additional spines not jjrocesses. Uropods 

a little longer than terminal abdominal segment. Outer 

branch about one-half as long as inner, which is two-thirds 

as long as peduncle. C. alabamohsis Stafford. 

BBB. Projjodus of first jjair of legs armed with three triangular processes. 

C. tridentata, sp. nov. 

AA. Propodus of first pair of legs not armed with triangular processes, but 
edged inside with spines. 

B. First pair of antenna\ with flagellum composed of eleven articles, 
extend one-third the length of the fifth article of the peduncle 
of the second antenna. Second antenna longer than the body; 
flagellum composed of about eighty-six articles. 

C. richardsonce Hay. 
BB. Fii-st pair of antennae, with flagellum composed of five articles, 
extend half the length of the peduncle of the .second antenna. 
Second pair of antennae "probably as long as body," flagellum com- 
posed of "at least forty segments." C . smithsii Ulrich. 

Coecidotea tridentata sp. nov. 

(Plate XV.) 

Size. The bod.\' without the antennae and uiopoda measures in length from 
9 mm. to 19 mm. and in width from 1% mm. to 3 mm. The length of the 
body is approximately five or s-'x t ines the width. This species is much larger 
than the others of this genus that have been describsd. From the descriptions 
I infer that 10 mm. is about the maximum of (*. styi/ia Packard, the largest 
member of the genus, while 3 mm. is supposed to be the maximum of C. 
smithsii Ulrich, the smallest. These figures suggest that the smallest mature 
C. triden'ata are about the size of some of the members of other species, but 
the laigpst indi\iduals are fully double that size. 

Color. The color is chalky white, the body wall being sufficiently clear 
to show the dark food canal within. Material .stored in alcohol appears very 
[lale yellowish gray. 

Stmchire. Head: Narrower than first thoracic segment. Wider than long. 
The cephalic margin narrower than the caudal, somewhat concave, and bear- 
ing the antennules and antennae, the bases of the latter appearing veiy heavy 
when compared with the size of the head. The antennule consists of the 
basal segments and a flagellum of from twelve to eighteen segments, the two 
parts of about equal length; the basal segment stoutest, a trifle longer than 
twice its width; second segment two-thirds as broad as basal and about 
.same length; third segment much smaller; five-eighths as broad and one- 
half as long as second. Its distal end bears the tai)ering flagellum. The an- 



Hungerford: A New Isopod. 177 

remui' arc r(>lati\ ely lar^r and consi.-^t of a ba.^al part of six segments and a 
flagelhim of from sixty to eifihty segments. Each of the first four segments 
of the basal ]iart is broader than h)ng. Taken together they are equal in 
length to th(> fifth segmtmt. wiiicii is a little shorter than the sixth, from which 
arises the many-segmented flagellum. The mandible bears a large three-seg- 
mented flattened jialp and two chitin-tipped processes, one a chisel-hke cutting 
edge and the other bearing from four to seven teeth. 

Thorax: The segments of the thorax are loosely articulated and their 
lateral margins are fringed with ver^• short, stout setae. All are broader than 
long. It bears seven pairs of legs, of which the first pair is subchelate. 

The first pair of legs is shorter than the others. In the males the propodus 
is very large and bears three well-develoi)ed processes, one at the base and 
two near the distal end. The ba^sal one is bifurcate in some and in others 
bears instead a strong seta. There are seven divisions to each limb, counting 
the clawlike terminal one. The propodus is the enlarged fifth division by this 
count. The limbs bear many strong setse and increase in length from the 
first to the last. 

Abdomen: The first two segments of the abdomen are short. The so-called 
third is nearl\- twice as long as wide and carries the uropods, each of wiuch 
consists of a basal part and two terminal branches. The uropods are longer 
than the abdominal segment which bears them, the relative length being 5:3. 
The basal segment is nearly as long as the last abdominal segment, the ratio 
about 6:7 in the males. The two branches are of very unequal length; the 
one female possessing inopods had this basal segment 1:3, the inner being 
much the larger. The relative lengths vary from 3:2 in the female to 4:1 in the 
male. The inner branch is to the peduncle as 3:4. There is considerable varia- 
tion in the comparative lengths of these parts. The second pleopod of the 
male, the first of the female and the third pleojiods of both sexes are unlike 
those figured by StafYord for C. alnbamensi.<<. 

Holotype, allotype and jiaratypes in alcohol. Kansas University' collection. 

Tiie females are smaller than the males and do not have as well- 
developad propodi. The sexual dimorphism appears not to have 
been recorded in the genus heretofore. Another point not mentioned 
in descriptions is the fact that the females possess the flattened 
brood pouches or oostegites at the base of the first four pairs of legs. 
Our specimens were obtained in June, and some of the females bear 
these plates. 

I asked Mr. Hoffmann, who gathered the material, to submit a few 
notes relating the circumstances of their collection. His notes in 
substance follow: 

The cistern frcni which these specimens were taken is about eight 
feet in diameter and nine feet deep. It contains a square brick 
filter, resting on the bottom in the center, which measures three or 
four feet square at base and tapers to its top, some four feet above, 
where it is just large enougli for a fotn'-inch casing, which extends 



178 The University Science Bulletin. 

above to within three feet of the top. This casing surrounds the 
pipe leading to the pump, and is open at the top. The water supply- 
ing the cistern is caught upon the roof of the house and conducted 
to the cistern by galvanized pipes three or four inches in diameter. 

On one occasion during a rain two isopods were observed by the 
lady of the house to be washed out of the elbow pipe leading from 
the gutter along the eaves of the house onto the sloping tin roof of 
the kitchen, thence into another gutter and down the pipe to the 
cistern. She concluded, therefore, that these animals, which she 
pumped up by way of the pitcher pump in the kitchen sink, were 
either "rained or had bred in the collection of wet leaves in the gut- 
ters of the house or in the elbow of the pipe leading from them." 

A number of specimens were taken alive in the water pumped 
from the cistern. One of these was placed in a specimen jar, three 
and one-half inches in diameter and three inches deep, where it 
lived in one and one-half inches of water from June 18 until July 26. 
The water was replenished from time to time with dirty pond water, 
containing many small organisms. 

Most of the specimens died within a few days. When several 
were placed together they seemed to take no notice of each other. 
The pleopods were observed to be in vibration as an individual 
made its way through the water. 

It is unfortunate that we were too busy to run any behavior exper- 
iments upon these most interesting forms. 

Note. — Through the kindness of the custodian of Crustacea I had the 
privilege of examining the Coecidotea material in the National Museum at 
Washington, D. C. One jar marked Coecidotea stygia contains eight vials; 
four of these contain large specimens which belong to the species I have de- 
scribed as new. It is interesting to note that they were collected at Topeka, 
Kan. They bear labels as follows : "Gift of E. A. Popenoe, Topeka, Kansas," and 
were taken "April 9, May 4, May 12, May 29, 1912." The other four vials con- 
tain material taken from "Graham's Spring, Lexington, Va., 1876"; Richard- 
son's Spring, Ky., W. P. Hay, Col.''; "Irvington, Ind., from wells W. P. 
Hay"; "Mammoth Cave, Ky., R. E. Call." 

The last four lots are much smaller specimens than the Kansas material. 
The material from Virginia, Richardson's Spring, Ky., and from Indiana, differs 
materially from the Kansas species. The species is broader than the Kansas 
crustacean, and the third pleopods are not only much broader comparatively, 
but are more truncate at the tip. 



1(S0 TriK UXIVERSITV SCIEN(,'K BULLETIN. 



PLATE XV. 

Ccecidotea tridentata (Crustacea). 

Fig. 1. First pleopod of female. 

Fig. 2. First antenna. 

Fig. 3. Mandible. 

Fig. 4. First pair pleopods of male. 

Fig. 5. Leg of female bearing oostegite. 

Fig. 6. Second pair pleopods of male. 

Fig. 7. Front leg of male. 

Fig. 8. Adult male. 

Fig. 9. Third pleopod of male. 

Fig. 10. Front leg of female. 

Fig. 11. Mature female carrying brood pouch. 



HrN(;KHFOHD: A New Tsopod. 



181 



PLATE XV. 




THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 7— October, 1922. 

(Whole Series, Vol. XXI^^ No. 7.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

Studies on Cicadella Hieroglyphica (Homoptera), 

Lucy M. Hackman. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



1/ 



TABLE OF CONTENTS. 

PAGE 

Introduction 189 

Life history notes 190 

Description of the species 190 

Hosts 192 

Hibernation 192 

Spring appearance 192 

Mating and oviposition 192 

Nymphs , 193 

Description of instars 194 

Adults 195 

Morphological studies 195 

General morphology of the abdomen 195 

Development of the male genitalia 196 

Development of the female genitalia: 198 

(187) 



I 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 7. 



Studies on CicadeUa hierogjyphica Say (Homoptera, 

Cicadellidae.) 

BY LUCY M. HACKMAN. 

Submitted to the department of entomology and to the graduate faculty of the University 
of Kansas in partial fulfillment of the requirements for the degree of master of arts, May IS, 
1922. 

INTRODUCTION. 

THE following notes on the life history of CicadeUa hieroglyphica 
consist of observations made from specimens in the field and in 
the laboratory. A growth of yomig willows along the Kansas river 
offered a splendid opportunity for the former, for there CicadeUa 
hieroglyphica may be found at all seasons in very large numbers. 
The laboratory observations were made from specimens collected at 
this place and reared on willow in the laboratory. The most satis- 
factory results M^ere obtained when the willow was planted in large 
glass rearing cages. The leaf hoppers could move about at will and 
were easily observed. 

Only a general description of the abdomen is given, for the chief 
concern of this paper is the genitalia. In tracing out the develop- 
ment of the genitalia in the male and female, the adult genitalia 
were used as a starting point. To trace the development, the various 
changes in the genitalia from one instar to another were studied. A 
study of the ventral surface of the eighth and ninth abdominal seg- 
ments was sufficient in the case of the female, for all three genital 
appendages are readily seen from a ventral view. But in the case 
of the male, where the two pairs of appendages are dorsal in position 
only, the development of the ventral ones could be traced by such a 
study. Therefore, particular attention was given to these dorsal or 
internal genitalia. For this purpose the pygofers were split open 

(189) 



190 The University Science Bulletin. 

along their dorsal surface, the overlying integument carefully re- 
moved, and the genitalia thus exposed. Great modifications in the 
genitalia occur within a single instar, and an attempt was made to 
give a rather detailed account of these modifications in the fifth 
instar. 

The writer wishes to express her appreciation to all who have as- 
sisted her in this work. Professor Hunter has always been most 
kind in helping in whatever way possible. Dr. Paul B. Lawson, 
under whose direction the work was done, has given freely of his 
time and experience. Kathleen Doering, Philip Readio and Robert 
Guntert are also deserving of thanks for their interest and assist- 
ance. 

LIFE HISTORY NOTES. 

The following are some of the references to this species: 

Tettigonia hieroglyphica Say, Jl. Acad. Nat. Sci. Phila., vi, p. 313, 1831. 

Tettigonia hieroglyphica Sig*> Ann. Soc. Ent. Fr., ser. 3, iii, p. 805, 1855. 

Tettigonia hieroglyphica G. and B., Hemip. Colo., p. 81, 1895. 

Tettigonia hieroglyphica Ball, Proc. la. Acad. Sci., viii, p. 51, 1901. 

Tettigoniella hieroglyphica Van D., Trans. San Diego Soc. Nat. Hist., ii, 
p. 52, 1914. 

Tettigoniella hieroglyphica Be L., Tenn. St. Bd. Ent., Bui. 17, p. 20, 1916. 

Cicadella hieroglyphica Van D., Cat. Hemip. N. A., p. 597, 1917. 

Cicadella hieroglyphica 01s., Bui. Am. Mus. Nat. Hist., xxxvii, p. 3, 1918. 

Cicadella hieroglyphica Lawson, Kan. Univ. Sci. Bui., xii, p. 85, 1920. 

DESCRIPTION OF THE SPECIES. 

The following is the original description: 

Tettigonia hieroglyphica. Dull rufous; head and scutel lineated; heme- 
lytra spotted. 

Inhabits Arkansas. 

Body obscurely dull rufous; head with a black dot at tii), above literate 
with black; thorax with a dusky jiosterior disk; scutel with black more or less 
curved lines; hemelytra obsoletelj' spotted, nervures being pale; beneath pale 
yellowish; pectus with large black spots; feet immaculate; tergum blue-black, 
edge yellow. Length to tij) of hemelytra one-fifth of an inch. 

Dr. P. B. Lawson, in his paper on the Cicadellidse of Kansas, gives 
the following description of the species: 

Form. Rather stout. Length, 6 to 7 mm. Vertex bluntly conical, wider 
than long. Pronotum nearly twice as wide as long, posterior angles broadly 
rounded, posterior margin medially emarginated. Elytra broad, but exceeding 
the abdomen. 

Coloi^. Varying from brick red to greenish and slaty blue. Black markings 
on vertex very strong and distinct, enclosing a light-colored T on basal half. 
Elytra with pale bands along the costal, claval and sutural margins. 



HaCKMAN : ClCADELLA HiEROGLYPHICA. , 191 

External genitalia. Fcmalo: Last ^•cntl•al sefinient about as wide as long, 
lateral margins triangularly luoduccd; pygofers long and narrow, equaling or 
slightlj' exceeding ovipositor, bearing a few stout hairs. Male: Last ventral 
segment less than twice as wide as long; plates long, broad at base, but 
tapering to long acute apices, margins fringed with short hairs; jiygofers long 
and narrow, equaling or exceeding plates and bearing stout hairs. 

Internal male genitalia. Styles short, distinctly bent in at point of attach- 
ment to connective by a large, heavily chitinized lobe, then curving outward 
and tapering gradually to bhmt apex, with an outwardly projecting process; 
connective slender, Y-shaped, stem of Y broadening to broad base; oedagus 
with pair of short processes extending dorsad from its point of attachment to 
connective, a long process leaving it dorsally from a point a little past its 
middle, and a similar longer one leaving it apically, the latter to the left of 
the former. These two processes are narrow and long, narrowest at the base, 
and widening to a point shortly before the apex, where they are the widest, 
the right one wider than the left one, and then tapering to the acute tips. A 
pair of somewhat narrow triangular chitinous processes extend from the base 
of the anal tube to the main body of the oedagus. 

//o.s"/.^. Taken abundantly on willows. 

The following variety occurs along with the typical forms: 

Cicadella hieroglyphica var. dolobrata (Ball). Its bibliography 

follows : 

Tettigonia hieroglyphica var. dolobrata Ball, Proc. la. Acad. Sci., p. 52, pi. 
3. fig. 2, 190L 

Tettigonia hieroglyphica var. dolobrata DeL., Tenn. St. Bd. Ent., Bui. 17, p. 
20, 1916. 

Cicadella hieroglyphica var. dolobrata Van D., Cat. Hemip. N. A., p. 597, 
1917. 

Cicadella hieroglyphica var. dolobrata 01s., Bui. Am. Mus. Nat. Hist., 
xxxviii, p. 3, 1918. 

Cicadella hieroglyphica var. dolobrata Lawson, Kan. Univ. Sci. Bui., xii, 
p. 86, 1920. 

Doctor Lawson describes this variety as follows: 

This is a smaller form than the preceding, appearing more robust. In color 
it is typically black, retaining a few of the lighter markings of the typical 
hieroglyphica on the front, vertex, pronotum and scutellum, and generally 
having the claval sutures light. 

Genitalia as in the preceding form. 

Distribution: Occurs along with the typical form. 

Hosts: Willows. 

DISTRIBUTION. 

Doctor Ball gives the following: "This species, as a whole, is 
very variable in size and color and recalls Oncometopia undata and 
lateralis in their red, green and black forms. The varieties readily 
fall into two series on structm-al characters. The first has hiero- 
glyphica and dolobrata as the extreme in darkening up. These 



192 The University Science Bulletin. 

forms are the only ones found in the Mississippi valley and as far 
west as central Kansas; they occur also in Texas, Arizona, and 
Mexico." 

Van Duzee reports it from Kansas, New Mexico, Texas, Illinois^ 
Missouri, Iowa, Nebraska, and Arizona. 

HOSTS. 

Cicadella hieroglyphica may be found on several hosts. Willows 
(Salix longi folia and Salix amygdaloides) are the most common of 
these, but it is frequently found on poplar {Populus monilifera). 
Occasionally it has been taken on the broad-leafed milkweed 
{Asclepias syriaca) and on giant ragweed (Ajnbrosia trifida) . 

HIBERNATION. 

During the winter the adults hide among the fallen leaves and 
rubbish on the ground, and appear very sluggish when disturbed. 
On mild, sunshiny days in January numbers have been observed 
sunning themselves upon the stems. 

SPRING APPEARANCE. 

About the middle of February, or when the willows are first be- 
ginning to bud, the greater number are to be found on the branches 
and stems of the willows. They are very gregarious, and often are 
so clustered together as to completely hide the stem. At this time 
of year they feed by sucking the sap from the stems, and give off 
honeydew in such quantities as to cause a noticeable spray. Upon 
close observation this honeydew is seen to be given off in a rapid 
succession of droplets from the anal tube. Several specimens were 
timed in the operation, and from fifteen to thirty drops were given 
off per minute. This continues for several hours at a time while the 
insect is feeding. Frequently this operation is accompanied by a 
spasmodic raising and lowering of the wings, movements of the 
abdomen, and stroking of the wings and abdomen by the long met- 
athoracic legs. 

MATING AND OVIPOSITION. 

(Plate XVII. figs. 1-4.) 

Early in April mating takes place. By this time the willow buds 
are beginning to unfold and oviposition begins. Numbers have been 
watched ovipositing, both in the field and in the laboratory, and the 
following observations made. 

The eggs are inserted in tlie tissues of the upper surface of the 
leaf just under the epidermis. In the act of ovipositing the female 



Hackmax: Cicadella Hi?:roglyphica. 193 

braces herself firmly, at times using her beak in addition to her 
legs for this purpose. In all eases observed she always worked head 
up. She first unsheaths her ovipositor, punctures the epidermis with 
its tip, and then inserts it to its full length. The flat surface of the 
ovipositor now rests parallel to the flat surface of the leaf, with its 
toothed edge pointing forward. Sawing the ovipositor back and 
forth she increases the size of the slit until it is large enough for the 
egg. The egg then passes between the valves of the ovipositor into 
the chamber prepared for it, and the ovipositor is withdrawn and 
sheathed. In a very few seconds the process is begun again. 

The time taken up in preparing the chamber and depositing the 
egg varied in several cases observed from two to five minutes, most 
of which time was spent in preparing the chamber. To cite a char- 
acteristic case, the whole operation occupied two and one-half 
minutes, the two minutes being spent in preparing the chamber and 
the half minute in placing the egg. 

The eggs may be laid singly or side by side in even rows. The 
largest number found in a single row was twenty-five and the 
largest number in a single leaf was thirty-five. The eggs in a hun- 
dred rows or masses were counted, and the average number per row 
was found to be seven. 

The effect of oviposition on the leaves is noticeable. The greater 
number of eggs are laid in leaves not fully developed, and the pres- 
ence of the eggs causes the growing leaf to become distorted and to 
curl around the eggs. However, in no case observed did oviposition 
kill the leaf. Eggs have been found in both willow and poplar 

leaves. 

NYMPHS. 

(Plate XVI, figs. 1-6.) 

Soon after oviposition, nymphs may be found feeding on the 
leaves. Eggs observed in the laboratory hatched in from eight days 
to two weeks. During their nymphal life these little leaf hoppers 
molt five times and become adult in a little over two months, or about 
the middle of June. By the middle of May the adults of the over- 
wintering generation are all dead. Shortly after becoming adult, the 
new adults mate and another generation is completed by the end of 
summer. This generation consists of the overwintering individuals. 
Xymphs af this generation have been found abundantly on giant 
ragweed and goldenrod, and in all probability the eggs for this gen- 
eration are laid in these hosts. 



194 The University Science Bulletin. 

DESCRIPTION OF THE INSTARS. 

For the description of the variout^ instars an attempt was made to 
select an average indivdual. It is possible to separate the males and 
females of the same instar by an examination of the ventral surface 
of the eighth and ninth abdominal segments. This is discussed later 
in greater detail under the development of the genitalia of each sex. 
Except for this, and a slight difference in size, the female being the 
larger, the two sexes are practically the same. 

Egg. 

(Plate XVII, fig. 1.) 

Length, 1.25 mm. Greatest width, .41. Subovoid in shape, somewhat pointed 
at one end; greenish yellow at first; just before hatching deeper yellow with 
dark e3'espots. 

First Instar. 

(Plate XVI, fig. 1.) 

Length, 2.65 mm. Width across eyes, .65 mm. Pale yellowish white, eyes 
black and prominent; anterior margin of head evenly rounded; two pairs of 
wing pads present, first pair short, barely covering base of second pair, caudal 
margin in form of an inverted W, with median projection extending farther 
caudad than lateral projections, exposed portion of second pair more than 
twice as long as first pair, caudal margin nearly straight, the segments marked 
off by light brownish bands. 

Second Instar. 

(Plate XVI, fig. 2.) 

Length, 3.15 mm. Width across eyes, .9 mm. Color same as in first instar; 
margin of head similar; wing pads have increased in length and width, but 
relative shape and position are the same; caudal margin of second pair bent 
slightly cephalad medially. 

Third Instar. 

(Plate XVI, fig. 3.) 

Length, 4.25 mm. Width across eyes, 1 mm. Color and shape of head un- 
changed; wing pads show decided change in length and width, lateral angles of 
first pair produced caudad and much longer than median projection; exposed 
length of second pair but little longer than first pair. 

Fourth Instar. 

(Plate XVI, fig. 4.) 

Length, 5.3 mm. Width across eyes, 1.35 mm. Color and shape of head 
practically the same as in preceding instars; lateral angles of first pair of wing 
pads now reach almost to the apex of the second pair; lateral angles of second 
pair have now extended farther caudad, making median indentation in caudal 
margin more pronoimced; do not extend beyond second segment of abdomen. 



Hackmax: Cicadella Hiekoglyphica. 195 

Fifth Instar. 

(Plate XVI, fig. .■).■) 
Lciititli, 0.4 nun. Width across eyes. 1.4 nun. Color the r^amc, head becom- 
ing!: slifrhtly more pointed, markings mon- distinct ; lateral angles of first pair of 
wing iiads still further jiroduced and are now about the same length as second 
pair; lateral angles of second pair also further produced caudad and now extend 
almost to fourth abdominal segment. 

Adults. 

(Plate XVI, fig. 0.) 

The seconci generation, or the overwintering one, consists of the 
typical reddish adults. Th? sexes can be easily distinguished one 
from the other by the darker color of the male abdomen as con- 
trasted with the lighter color of the female's. These produce the 
slaty-gray individuals of the summer generation. The black form, 
the variety dolobrata, is also present at this season. However, only 
males of this form have been found. They mate with the slaty-gray 
individuals, which are for the most part females, although males of 
this type are numerous. Very evidently, the dark forms are di- 
morphic males of the summer generation. A single dark male was 
taken April 1 with the overwintering red forms, but whether this is 
one of the summer forms which has overwintered or one of the win- 
tering generation which differs in color from the rest of the gener- 
ation has not been determined. 

MORPHOLOGICAL STUDIES. 
GENERAL DESCRIPTION OF THE ABDOMEN. 

(Plate XVIII, figs. 1-3; (5-9.) 

The abdomen is joined broadly to the metathorax. It continues 
at the same width for about two-thirds of its length, and from there 
tapers to a somewhat pointed apex. In a general cross section it is 
semicircular in outline. The sternite and the pleurites, ventral in 
position, form the straight part of the semicircle, while the curved, 
dorsal tergite forms the circular part. Eleven segments can be ac- 
counted for. In the male, six of these are represented by complete 
sternites. pleurites and tergites, but in the female only five are so 
represented. In the first two segments only the sterna and terga are 
present, the pleura being represented by pleural membranes. Seg- 
ments three to eight in tiie male and three to seven in the female are 
typical. The terminal segments in both sexes are modified. These 
modifications are discussed under the heading "external genitalia" 
in the description of the species. In addition to what is given there, 
I sliould like to add that the ventral valve of the adult is present, 
but concealed by the last ventral segment. 



196 The University Science Bulletin. 

DEVELOPMENT OF MALE GENITALIA. 

(Plate XIX, figs. 2, 4, 6, 8, 10, 12; Plate XX, figs. 1-18.) 

The male genitalia, consisting of three pairs of valves arise from 
a genital area on the ninth abdominal segment. In the first, second 
and third instars there are two pairs of valves present. These de- 
velop in small chitinous pockets, which are attached at the caudal 
edge of the genital area with their apices directed caudad. The 
genital area increases slightly in size in each successive instar, as do 
the pockets. The pockets are placed one upon the other, the ven- 
trally placed pocket producing the plates of the adult and the more 
dorsal pair the oedagus. The ventral pair is about twice as long as 
the dorsal. Both taper caudad, and are somewhat rounded at the 
apex. Each pair is divided into its right and left valve by a chit- 
inous median partition. 

In the fourth instar there is, as in the former instars, a noticeable 
increase in size, both of the genital area and of the ventral and dor- 
sal pockets. In addition to these structures, there is now present a 
pair of small lateral pockets located at the lateral margin of the 
genital area, and extending caudad as far as the bases of the dorsal 
pockets. In these lateral pockets develop the styles of the adult 
genitalia. In the fifth instar there is no great change in the relati^'e 
position and shape of the pockets. There is, of course, a natural 
increase in the size of all three pockets. 

In the fifth instar the greatest changes in- the developing genital 
appendages take place. The three pairs of valves may be traced 
through several distinct phases of development by a study of the 
soft, white integument which can be drawn from the chitinous pock- 
ets. Details of five particular phases might be mentioned. (Plate 
XX, figs. 6-18.) 

The ventral plates which develop in the ventral pockets show 
very little change throughout the successive phases of this instar. 

The styles which develop in the lateral pockets are present in all 
five phases and show a gradual increase in size through the phases. 
In the fourth phase they first show their permanent attachment to 
the ventral plates, which is more apparent in the fifth phase and in 
the adult. They are attached on their outer edges near the base of 
the plate. 

The developing oedagus or the integument drawn from the dorsril 
pockets shows the most remarkable alterations. In phase 1 the 
cedagus consists of two valves placed parallel to the ventral plates. 



Hackman: Cicadella Hieroglyphica. 197 

In phase 1, ventral aspect, the valves placed side by side are com- 
paratively narrow at their bases, widest at a point a little before the 
middle, and then tapered to somewhat pointed apices. Dorsally 
they do not appear to extend as far cephalad as they do ventrally, 
nor are thej^ divided into right and left valves except for a short 
distance apically, at which point they are widely separated. 

In phase 2, viewed ventrally, the two valves have become longer 
and slightly narrower than in the preceding phase, and instead of 
their former relationship, side by side, the right valve at the base is 
now assmning a ventral position with respect to the left valve. Also, 
the flat surfaces of the valves, instead of being parallel to the ventral 
plates, are now assuming a perpendicular position. Between the 
valves, at a point a little beyond the middle, a short, slender finger- 
like process is visible. Viewed dorsally, this process appears be- 
tween the two valves at the point of wide separation in phase 1 and 
projects for a short distance caudad. It is an evagination of the 
integument of the valves. 

In phase 3, ventral aspect, the two valves have continued to in- 
crease in length. The right valve is distinctly folded over the left 
valve at the base. A lateral aspect shows plainly the relative posi- 
tion of the median process and the valves. In phase 3, dorsal aspect, 
there is little change in the valves except an increase in length. The 
median process also shows an increase in length and width. 

In phase 4, ventral view, the valves again are longer than in the 
preceding phase, and the median process also shows a distinct in- 
crease in length. The folding of the right valve over the left valve 
is more complete, and the bases of the two valves are farther apart 
than in the preceding phase, the left valve having moved caudad. 

In phase 5 the oedagus appears very similar to the adult oedagus. 
Here the valves have become very much longer than in phase 4. 
The median process has increased greatly in length. 

During the development of the valves their apices have main- 
tained their relative length in respect to the ventral plates. The in- 
crease in length has been taken up by increased dorsal curvature. 

For a description of the adult male internal genitalia, reference 
may be made to the description of the species. 



198 The University Science Bulletin. 

DEVELOPMENT OF THE FEMALE GENITALIA. 

Ovipositor of Adult. 

(Plate XVIII. figs. 4-6.) 

In the female the genitaha also consist of three pairs of append- 
ages. They are the ventral and dorsal pairs, which make up the 
ovipositor proper, and the lateral pair, within which the ovipositor 
is folded. The ventral pair arise from the eighth sternum, and the 
dorsal and lateral pairs arise from the ninth sternum, the dorsal 
pair from its cephalic margin and the lateral pair from its caudal 
margin. 

The ventral valves arising from the eighth sternum are two long, 
slender valves, tapering caudad and sharply pointed at the apices. 
They are ventral in position and inclose the dorsal valves. The 
ventral and dorsal valves of each side are joined one to the other 
by a tongue-and-groove arrangement, the tongue being on the dor- 
sal valve and the groove on the ventral. 

The dorsal valves are innermost of the three pairs and are dorsal 
in location. They consist of a narrow, curved, rodlike base, and a 
broader, flattened apical part which bears teeth along its dorsal 
edge. The apex is sharply pointed and bears very fine teeth on 
both dorsal and ventral edges. There are from seventeen to nine- 
teen large teeth on the dorsal edge. 

The lateral valves are the outermost of the three and enfold the 
other two pairs. They are broad, flat, and somewhat concave on 
their inner surfaces. They are not capable of extension as are the 
other two pairs, for in addition to being attached basally to the 
ninth sternum, they are also attached to the pygofers for about one- 
half their length. Their apical half is free. 

Nymphal Genital Appendages. 

(Plate XIX, figs. 1, 3, 5, 7, 9, 11.) 

In the first and second instars two pairs of very small chitinous 
pockets may be seen, one projecting caudad from the eighth sternum 
and the other from the ninth. In the first pair develop the ventral 
valves of the adult ovipositor, and in the second the dorsal valves. 
In these two instars the apices of the ventral pockets are at a dis- 
tance from the bases of the dorsal pockets. The dorsal pockets, 
extending caudad for about one-third the length of the ninth seg- 
ment, are also short. 

In the third instar the ventral pockets have increased in length 
and width. They are still short and broad. Their broadly rounded 



Hackman: Cicadella Hieroglyphica. 199 

apices overlap the bases uf the dorsal pockets. The narrower, 
longer dorsal pockets extend caudad for about one-half the length 
of the ninth segment. In this instar the lateral pockets first appear. 
They lie laterad of the dorsal pockets, are slightly curved and some- 
what narrower and shorter than these. 

In the fourth instar the ventral pockets are longer than in the 
preceding instar, and are now more than two-thirds the length 
of the dorsal pockets. The ventral, still the longer of the two pairs, 
are about the same distance from the tip of the ninth segment. 
They are beginning to be enfolded by the lateral pockets, which 
have broadened and are now somewhat concave on the inner sur- 
face. The lateral pockets are longer than the ventral pockets, but 
shorter than the dorsal. 

In the fifth instar the three pairs of pockets are rather darkly 
chitinized. The ventral pockets are broad at the base and taper 
gradually to narrowly rounded apices. They are the shortest of 
the three pairs, though nearly as long as the dorsal pair. Only the 
extreme apices of the dorsal pockets are visible, their basal portion 
being hidden by the broader ventral pockets. They are still longer 
than the ventral pockets, but are now exceeded in length by the 
lateral pockets. These are somewhat spoonlike and have more com- 
pletely enfolded the dorsal pockets. 



200 - The University Science Bulletin. 



PLATE XVI. 

1. First instar. 

2. Second instar. 

3. Third instar. 

4. Fourth instar. 

5. Fifth instar. 

6. Adult. 



Hackman: Cicadklla Hip:roglyphica. 



201 



PLATE XVI. 






■r^.vv".-.-,— ^ 


k. 


.^..,.,J 


p'.^"'-"'.'"'"^' 




1 


1 


t 


J 




T*^ 


1^..., 


J 






C^ 


"^"Z^ 


L, 


.,.,.■, ^,.V.^-'.,'».t.'<>.'^ 


C , 


.....„..i 


r 5 


\ -/ 






^ * 



6 



202 The University Science Bulletin. 



PLATE XYH. 

1. Egg enlarged. 

2. Leaf with two egg masses. 

3. Leaf showing effect of ovii)ositioii. 

4. Twig showing effect of oviposition. 



Hackman: Ckadkli.a Hierchjlyphica. 



203 



PLATE XVII. 




204 The University Science Bulletin. 



PLATE XVIII. 

1. Abdomen of adult female. 

2. First sternite of female abdomen. 

3. Second sternite of female abdomen. 

4. Ventral valve of ovipositor attached to eighth sternum. 

5. Doi-sal valves (upper) and lateral valves (lower), showing at'acl.int iit 
ninth sternum. 

6. Terminal segments of female abdomen, showing ovipositor. 

7. First sternite of male abdomen. 

8. Second sternite of male abdomen. 

9. Abdomen of adult male. 

10. Cephalic view of main body of cedagus. 

11. Caudal view of main body of oedagus. 

12. CEdagus, showing attachment of ejaculatory duct. 

13. Styles and connective. 

14. Ventral valves, dorsal aspect, showing attachment of styles. 

15. Male genitalia. 



HacKMAN: ('l(AI)KLLA II 1 KKOdl.VI' 1 1 ICA. 



205 



PLATE XVIII. 



%.=^ 




206 The University Science Bulletin. 



PLATE XIX. 

Terminal. Segments of Abdomen; 

1. Female, first instar. 

2. Male, first instar. 

3. Female, second instar. 

4. Male, second instar. 

5. Female, third instar. 

6. Male, third instar. 

7. Female, fourth instar. 

8. Male, fourth instar. 

9. Female, fifth instar. 

10. Male, fifth instar. 

11. Female adult. 

12. Male adult. 



Hackman: Cicadella Hieroglyphica. 



207 



PLATE XIX. 




208 



The University Science Bulletin. 



PLATE XX. 



Development of Male Internal Genitall\: 

1. First instar. 

2. Second instar. 

3. Third instar. 

4. Fourth instar. 

5. Fifth instar. 

6. Fifth instar, phase 1, dorsal view. 

7. Fifth instar, phase 1, ventral view. 

8. Fifth instar, phase 2, dorsal view. 

9. Fifth instar, phase 2, ventral view. 

10. Fifth instar, phase 3, dorsal view. 

11. Fifth instar, phase 3, ventral view. 

12. Fifth instar, phase 3, lateral (right) view of oedagus. 

13. Fifth instar, phase 3, lateral (left) view of cedagus. 

14. Fifth instar, phase 4, ventral view of cedagus. 

15. Fifth instar, phase 4, lateral (right) view of oedagus. 

16. Fifth instar, phase 4, lateral (left) view of cedagus. 

17. Fifth instar, phase 4, lateral (right) view of oedagus. 

18. Fifth instar, phase 4, relative position of plates, styles and oedagus. 



Hackman: Cicadella Hieroolvphka, 



209 



PLATE XX. 



3 




THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 8— October, 1922. 

(Whole Series. Vol. XXIV, No. 8.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 
Ovipositors of Cicadellide (Homoptera) P. A. Readio. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE, KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



TABLE OF CONTENTS. 

PAGE 

Introduction 217 

The abdomen of the female 218 

The ovipositor 220 

Taxonomic use of the ovipositor 222 

Descriptions of the ovipositors of the genera and species 224 

Conclusions 264 

Plates 267 

Bibliography 265 

Index 293 

(215) 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 8. 



The Ovipositors of the CicadelUdae (Homoptera) . 

By PHILIP A. READIO. 

.•^iihniitted to the department of entomology and to the siaduHte faculty of the University 
of Kansas in partial fulfillment of the requirements for the degree of master of science, 
May 1.'), 1922. 

INTRODUCTION. 

THE primary purpose of this paper is to determine whether or 
not the ovipositors of the Cicadelhclse possess characters of 
value in classification, and if so, of how much value these characters 
are and how accessible they are to the general worker. To Prof. 
Paul B. Lawson belongs the credit for suggesting the paper. In the 
same group he has recently found that the male genitalia are of 
much value in classification, and the possibility of equal value in 
the female genitalia occurred to him. Miss Itasca Hilsman, work- 
ing in the closely related family, Cicadidse, found that the ovi- 
positors in this group "afford constant and ready characters which 
may at times be of decided value to him (the specialist) in the de- 
termination of closely related species." Hence it was natural to ex- 
pect that equally favorable results might be obtained from a study 
of the ovipositors of the Cicadellidse. As an introduction to the 
taxonomic part of the paper, a morphological study of the abdomen 
of the female and the structure of the ovipositor was made. 

In addition to the reasons already stated, there are certain gen- 
eral considerations which would lead one to expect such an investi- 
gation as this one to be fruitful. It is constant characters that the 
taxonomist is in search of — characters that are sufficiently definite 
to separate closely related species, and yet are present in tlie entire 
range of the species. Because of their internal position, genitalia 
are more likely to be constant than external structures, which may 
vary with differences of environment. The constancy of the use to 
which genitalia are put also makes for the permanency of their 
structure and their usefulness as taxonomic characters. 



(217) 



218 The University Science Bulletin. 

The material necessary for this study was obtained from the 
duplicate collection of the University of Kansas entomological mu- 
seum. A representative number of ovipositors from each of the 
subfamilies, excepting the Paropinse, was examined, and an attempt 
made to examine as many different genera and as many species in 
each genus as possible. In all, representatives of forty-eight genera 
and ninety-two species were examined and figured. 

The writer wishes to express to Prof. S. J. Hunter his appreciation 
of the sanction given to this work, and of the necessary time and 
materials so kindly placed at his disposal. To Prof. P. B. Lawson 
many thanks are due, both for the conception of the nature of the 
problem and for the direction of the work to its completion. It 
was under the able direction of Prof. P. W. Claassen, of Cornell 
University, that the photography was done. To Prof. H. B. Hunger- 
ford, Miss Lucy Hackman, Miss Kathleen Doering and Mr. Robert 
Guntert the writer wishes to express his appreciation for their kind 
help, suggestions and criticisms. 

THE ABDOMEN OF THE FEMALE. 

(PI. XXI, figs. 1-3.) 

For a study of the abdomen of the female leaf hopper, dried speci- 
mens of Oncoinetopia lateralis (Fabricius) were used. These were 
soaked in caustic potash, ten per cent, for twenty-four hours, and 
drawings and descriptions made from the cleared specimens. 

The abdomen joins the thorax broadly, bulges slightly in the mid- 
dle, and, from a point a little past the middle, tapers both in width 
and in height to a blunt, caudal point. A cross section has a general 
semicircular outline. The tergum is arched, appears dorsally and 
laterally, and forms the rounded portion of the semicircle, while the 
pleura and sternum are flat, and form the fiat, ventral portion. The 
lateral edges of the sternum are bent slightly dorsad at their union 
with the pleura. The pleura and tergum unite at a distinct but 
slightly rounded angle. 

Eleven segments can be accounted for in the abdomen. Of these, 
segments one and two lack distinct pleura, and segments eight to 
eleven are variously modified as described. 

The tergum of the first segment is partly membranous and partly 
chitinized. Cephalad, at its junction with the metathorax, it is en- 
tirely membranous. Caudad, at its junction with the second ab- 
dominal segment, is a narrow, linear, chitinized sclerite which bears 
laterally two inwardly projecting processes along its cephalic border. 



Readio: Ovipositors of Cicadellid.f.. 219 

Between the inwardly projecting processes is a round, small, chi- 
tinized piece. The pleura are entirely membranous, and in the 
pleural membranes, opening ventrally, are the spiracles of the first 
segment. These are larger than those of the other abdominal seg- 
ments. The sternum consists of a curved, lightly chitinized caudal 
sclerite extending for nearly the entire width of the segment, and of 
a membranous cephalic portion connecting with the metatliorax. 
The chitinized portion has along its cephalic border two pointed 
projections which point mesad. 

The tergite of the second segment consists of a simple, linear, 
chitinized sclerite which extends nearly the entire width of the seg- 
ment. It is about one-third the length of the third tergite. The 
pleura are represented only by pleural membranes in which the 
spiracles of this segment are located. The spiracles open dorsally 
and appear to be in the membrane laterad of the second tergite. 
The sternum of segment two is a curved, chitinous piece extending 
the entire width of the segment. 

Segments three, four, five and six have distinct and complete 
terga, sterna and pleura. The terga are arched and form the dorsal 
and lateral surfaces. The ventral surface is formed by the pleura, 
which are subrectangular and longer than wide, and by the sterna, 
which are also subrectangular, but wider than long. Each pleuron 
bears a spiracle in a cephalomesal position. The fourth segment 
is slightly longer than the third, and the fifth and sixth are about the 
same length and longer than the fourth. 

Segment seven is also complete. The tergum and pleura are the 
same as in the preceding segments. The sternum forms the sub- 
genital plate and is commonly called the last ventral segment. Its 
lateral margins converge caudad and its caudal border bears a broad 
but shallow indentation along its mesal half. Dorsad of the sternum 
of segment seven is an invaginated pocket in which the bases of the 
valves of the ovipositor are located. Its ventral side is formed by 
the apical portion of the seventh sternum and a membrane arising 
from the dorsal surface of this sclerite and extending cephalad to 
its base. The dorsal side of the pocket is formed by a membrane 
which is continuous with the membrane of the ventral side, and 
which extends caudad to the eighth sterna with which it connects. 
The greater part of this dorsal membrane is strongly chitinized, but 
medially it is apparently divided by a narrow, clear line which ex- 
pands apically into the entirely membranous apex connecting with 



220 The University Science Bulletin. 

the eighth sterna. Laterally this pocket is bounded by the continu- 
ation of the dorsal membrane to the seventh and eighth pleura. The 
anterior portions of these lateral membranes are strongly chitinized 
and curve mesad ventrally. 

The terminal segments of the female abdomen bear the external 
organs of reproduction and are modified for this purpose. Segment 
eight i^ represented externally only by the tergum and the pleura. 
The tergum is essentially the same as in the preceding segments, 
but narrows slightly caudad. The pleura narrow caudad and are 
triangular in shape. They include the ventral cephalic portion of 
the ninth segment between them and bear the last pair of spiracles. 
Externally segment nine consists only of a tergum, commonly called 
the pygofer, which almost completely encircles the abdomen, leav- 
ing only a ventral groove in w'hich the ovipositor is folded. Ven- 
trally it extends cephalad, ending in two obtuse points between the 
eighth pleurites. Tlie caudal portion encircles nearly the whole seg- 
ment and tapers towards its caudal end. 

The anal tube is composed of segments ten and eleven. Segment 
ten arises from the narrowed apex of segment nine and is short and 
tubular. It connects by a distinct intersegmental membrane with 
segment eleven, which is also tubular and can be telescoped within 
segment ten. Segment eleven bears the telson, the extreme apical 
portion of the abdomen. Along the dorsal edge of the telson is the 
anus, an opening running for its entire length and guarded by hairs. 

There are eight spiracles in the abdomen of the female located in 
the first eight segments as described. 

Along the ventral side of the abdomen there occur many short, 
fine hairs, which in all cases point backward. The hairs on the ninth 
segment, or pygofer, are somewhat longer than those on the other 
parts of the abdomen. 

THE OVIPOSITOR. 

(PL XXI, figs. 4-9.) 

This description was made from the ovipositor of Oncometopia 
lateralis (Fabricius). However, in the many other species of leaf 
hoppers examined, the fundamental structure of the ovipositor was 
lound to be the same as for the species here described. 

The ovipositor of the leaf hopper is fitted for sawing slits in plant 
tissues and for placing eggs within these slits. When not in use it 
lies in the groove in the ventral surface of the ninth segment. The 
ovipositor consists of three pairs of valves or gonapophyses. The 



Readio: OviPCSiTORs of Cicadellid.e. 221 

inner two pairs alone t'uiictioii in sawing slits and placing eggs, and 
the outer pair act as guards within which the two inner pairs lie 
when not in use. In this discussion the valves are numbered I, II 
and 111, according to their attachment, the most cephalic in attach- 
ment being numbered I. 

Valve I is the outer and more ventral of the two inner pairs of 
\'alves. Its chief attachment is to the eighth sternum, which is 
divided and appears as two subrectangular, heavily chitinized 
sclerites, for the most part hidden in the pocket above the seventh 
sternum. Near its base, a narrow, curved rod leaves its dorsal mar- 
gin and attaches basally to a small, triangular sclerite on the dorsal 
side of the cephalic end of the pygofer. This small triangular scle- 
rite is possibly the ninth pleurite. Upon attempting to separate the 
narrow, rodlike portion from the rest of the valve, a featherlike 
structure, consisting of a middle portion formed by the continua- 
tion of the attaching rod and membranous side plates, can be drawn 
out. However, except basally, this featherlike structure is en- 
tirely incorporated into the main part of the valve, both structurally 
and functionally. The main part of the valve is relatively broad 
basally, narrows slightly beyond the base, broadens again towards 
the middle, and narrows to a sharp, apical point. A heavily chitin- 
ized rod strengthens the ventral portion of this valve. The rod 
is comparatively large basally, narrows in the middle portion, and 
is absent in the apical third. Along the entire dorsal border and 
along the ventral apical border are diagonal rows of scalelike proc- 
esses. Along its mesal surface, just dorsad of the chitinized rod, is 
a mushroom-shaped groove into which a complementary tongue on 
the outer surface of valve II fits. This permits independent sliding 
of the two valves, but not complete separation. Basally the ven- 
tral edges of this vah-e turn mesad, and on the dorsal side of this in- 
turned portion is a less heavily chitinized projection. This projec- 
tion in the one valve bears a tongue, and in the other a groove, unit- 
ing the two basally. For the greater part of their length, however, 
they are independent. 

Valve II is the inner and the more dorsal of the two inner pairs 
of valves. It is attached to tlie cephalic end of a small sclerite, 
which is attached to the pygofer. This sclerite is ventral in posi- 
tion and possibly represents the ninth sternite. This valve is narrow 
and rodlike at the base, but for its greater length is broad and flat. 
It also possesses a chitinized strengthening rod whicli extends nearly 
to its apex. On the outer surface of the valve, just ventrad of the 
rod, is a tongue which fits into the groove in valve I. The dorsal 



222 The University Science Bulletin. 

edge usually bears teeth, which may extend for its entire length, be 
confined to the apical half, or appear only at the tip. The teeth 
are of various shapes in the different species, and may or may not 
in turn bear secondary teeth. The pointed tip is usually notched 
with small teeth, and these may appear on either the dorsal or ven- 
tral or both sides. From the interior of the valve certain ductlike 
structures lead to the dorsal and apical teeth. The function of these 
ducts is unknown. This pair of valves is usually united one to the 
other basally, sometimes by a heavy chitinized connection, some- 
times only by a membrane, but not by a tongue and groove. 

Valve III is the most caudal in attachment, being attached to the 
caudal end of the same sclerite to which valve II is attached. There 
is also a membranous connection between this valve and the ventral 
side of the pygofer for about h;ilf the length of the valve. The 
proximal half of the valve is narrow and the apical half broad and 
spoonlike, being somewhat concave on its inner surface. It is 
bluntly rounded at the apex. This pair of valves, between which 
there is no connection, forms a protecting sheath in which the ovi- 
positor is completely encased when not in use. 

Oviposition of Cicadella hieroglyphica (Say) has been observed 
and the use of the ovipositor noted. Valve III remains in its posi- 
tion in the groove of segment nine and takes no part in the opera- 
tion. Valves I and II, closely appressed and appearing as a single 
structure, are extruded from their resting place and form the func- 
tional part of the ovipositor. The ovipositor is first held vertically 
and its tip inserted under the epidermis of the leaf. It is then 
pushed in to nearly its full length, and now is in a horizontal posi- 
tion, the flat surface of the ovipositor being parallel with the flat 
surface of the leaf. The teeth on the dorsal edge of the ovipositor 
are now turned cephalad. The ovipositor is pushed backward and 
forward with a sawlike motion until the chamber is large enough for 
the reception of the egg. The egg passes out between the valves of 
the ovipositor and into the chamber, and when in place, the oviposi- 
tor is withdrawn and folded into its resting place. 

* 

TAXONOMIC USE OF THE OVIPOSITOR. 

In the search for characters of taxonomic value in the ovipositor, 
valve II was at once hit upon as being the structure most likely to 
possess useful characters. It varies in different species in regard to 
size, shape, number and shape of primary teeth, number and shape 
of the secondary teeth borne by the primary teeth, characters of the 
tip, and in the number and arrangement of the ducts. That these 



Readio: Ovipositors of Cicadp:llid^. 223 

characters are constant within (he species has been proven by ex- 
amining a wide range of in(h\-i(hials within the species and finding 
that the \-ariation was negligible. C'onst'ciuently, for the taxononiic 
part of this paper, valve II is used exclusively. 

The technique used in mounting valve II for study is simple. The 
tip of the abdomen bearing the oN'ipositor is broken off from the 
dried specimen, soaked in ten i:>er cent caustic potash for twenty- 
four hours, washed in water for a few minutes, valve II and other de- 
sired parts dissected out under a binocular microscope, dehydrated 
in ninety-five per cent alcohol for five minutes, cleared in xylol for 
five minutes, and mounted in balsam on a microscope slide. It is 
well to give the slide and the specimen from which the slide w^as 
made a corresponding number, so that any necessary checks may 
be made. This method gives a permanent mount, which may be 
studied at the convenience of the worker. 

Several methods of figuring the ovipositor were used. The first 
was that of drawing the desired valve free-hand with the aid of a 
micrometer eyepiece divided into squares, which correspond to 
squares on the drawing paper. This is a satisfactory method, but 
somew^hat more laborious than the methods later used. The second 
was to draw with the aid of the camera lucida. The particular 
equipment available did not give satisfactory results at all times, 
but there is no doubt but what this method could be used with 
satisfaction. The greatest degree of success was obtained by the 
use of the Edinger drawing apparatus. This apparatus projects the 
desired image upon the drawing paper, and the figuring consists only 
of tracing in the image. Photography w'as tried and found to be a 
very successful means of reproducing the desired image. A camera 
fitted for ordinary scientific work was used. It w^as turned to a ver- 
tical position and a lens board fitted with a black, light-proof sleeve 
was inserted in the front lens-board holder. The sleeve was lowered 
over the ocular of a compound microscope, and focusing for desired 
size and definition of image accomplished by a combination of 
microscope and camera adjustments. Various substitutions in ob- 
jectives and oculars were necessary to meet all conditions. Illumi- 
nation was furnished by an ordinary substage light, and exposures 
were made by the switching off and on of this lamp. Undoubtedly 
much better results could be obtained with the aid of special photo- 
micrographic equipment. 

Descriptions of the ovipositors of the generji and species of the 
various subfamilies follow: 



224 The University Science Bulletin. 

Subfamily BYTHOSOPIN.^ (Dohrn). 
Agalliopsis novella (Say). 

(PI. XXII, fig. 2.: pi. XXV, fig. 1.) 

Length, 1.1 mm.; greatest width, 0.11 mm. Apical toothed half only 
slightly wider than basal half; distinctly curved, tip rather gradually narrowed, 
extreme apex somewhat pointed, chitinization light; strengthening rod ex- 
tends caudad as far as twenty-first or twenty-second dorsal tooth from apex. 
Toothed area on doi-sal edge occupies slightly less than the apical half, teeth 
sixty-three to sixty-five in number, very small, angular, pointed, rather regu- 
lar in size, shape and spacing; a few double teeth present, no secondary teeth; 
the tip is notched on both edges with small teeth, practically continuous 
around the tip, twenty to twenty-two on \'entral edge; area of ducts in- 
conspicuous, a few ducts faintly visible for their entire length, but for the 
most part are invisible except for their apices and circular openings, open 
along the ventral apical edge (seven), along the dorsal edge of the toothed 
area, and in the basal region; the two valves of the pair are joined one to 
the other by a narrow, elongate, chitinized connection present on the dorsal 
edge of the basal region at a point about two-fifths the length. 

Aceratagallia uhleri (Van Duzee). 

(PI. XXV, fig. 2.) 

Length. 1 mm.; greatest width, .07 mm. Apical half very little if any 
wider than basal half; .distinctly curved, tip gradually narrowed, extreme 
apex rather pointed, chitinization moderately light; strengthening rod ex- 
tends caudad as far as twenty-second dorsal tooth from apex. Toothed area 
on dorsal edge occupies the apical half of the length; teeth about ninety 
in number, very small, angular, pjointed, fairly regular as to size, shape and 
spacing, larger apically; bear no secondary teeth; the tip is notched with small 
teeth on both edges, continuous around the tip, about twenty-two on the 
^•entral edge; area of ducts conspicuous, ducts easily visible for entire length; 
open along ventral apical' edge (five to six), along dorsal edge of toothed 
area, and in basal region; the two valves of the pair are joined one to the 
other by an elongate, naiTOW, chitinized connection present on the dorsal edge 
at a point about one-third the length. 

This ovipositor is similar to that of Agalliopsis novella (Say) in 
general appearance. 

Genus Idiocerus Lewis. 

The ovipositors of seven species of this genus have been examined 
and found to be generally similar. In each case the ovipositor is 
about the same width for the entire length and is only slightly 
curved. Examples of light, medium and heavy chitinization are 
found. The valve may be toothed along its dorsal edge from a 
fourth to a little more than a half its length apically. The teeth 
in the specimens examined are ten to thirty in number, medium to 
large in size, rounded, evenly spaced, and may or may not bear 
secondary teeth. The ducts may appear ductlike, granular, or be 



Readio: Ovipositors of Cicadellid.e. 225 

invisible. The apex may or may not bear small teeth along one 
or both edges. A ohitinous connection is present on the dorsal edge 
of the basal area in some species. 

Idioccrus ncrvatus Van Duzee. 

(PL XXV, fig. 9.) 

Length, 1.4 nun.; greatest width, 0.12 nun. Apical portion slightly wider 
than basal portion; shghtly curved, tip greatly narrowed with the extreme 
apex bluntly rounded, veiy lightly chitinized; strengthening rod extends 
caudad as far a^; eleventh doi-sal tooth. Toothed area on dorsal edge occupies 
the apical two-fifths of the length; teeth fourteen to sixteen in number, large, 
long, rounded, regular, evenly spaced, though somewhat farther apart basally; 
bear no secondary teeth, though the margin is irregularly roughened; the tij) 
is notched on' both edges with small, forward-pointing teeth, eight appear 
ventrally, one to three dorsally; area of ducts inconspicuous, circular openings 
alone visible; of these four open ventrally, three apically, and many dorsally. 
There is no evidence of a chitinous connection. 

Idiocerus pallidus Fitch. . 

(PI. XXII, fig. 1; p!. XXV, fig. 5.) 

Length, 2.4 mm.; greatest witlth, 0.25 mm. About the same width for 
entire length; slightly curved, tip bluntly rounded, heavily chitinized; strength- 
ening rod extends caudad as far as ninth dorsal tooth. Toothed area on 
doi-sal edge occupies a little more than the apical third; teeth fourteen in 
number, large, rounded, regular, evenly spaced except basally, where they are 
farther apart; secondary teeth only on last eight primary teeth, these very 
small and indistinct, especially on those farthest from the tip. where they can 
be seen only with the aid of high-power magnification; the tip is notched with 
small, regular teeth on both dorsal and ventral edges; area of ducts con- 
spicuous, ducts elongate, large, straight; six open along ventral apical edge, 
one in extreme apex, and many along the dorsal edge. A rather indistinct 
chitinous connection is present on the doi-sal edge of the basal area and joins 
the two valves of the pair. 

Idiocerus diizeei Provancher. 

(PI. XXV, fig. 6.) 

Length. 2.6 mm.; greatest width, 0.34 mm. Apical portion slightly wider 
than basal portion; slightly curved, tip ends in somewhat rounded, obtuse 
point, heavily chitinized; strengthening rod extends caudad as far as sixth 
dorsal tooth. Toothed area on dorsal edge occupies a, httle less than the 
apical half, teeth nine to ten in number, large, somewhat angular, regular, 
evenly spaced, but larger and farther apart basally, teeth one and two being 
veiy large, rounded, and heavily chitinized; all the primary teeth except 
teeth one. two and three are notched with small secondary teeth; tip is 
notched with many small, regular teeth on both dorsal and ventral edges; 
ducts conspicuous, rather small, straight, open by circular openings; six open 
along the ventral apical edge, one in extreme apex, and many along the 
doi-sal edge; there is no well-defined chitinized connection between the two 
valves of the pair. 



226 The University Science Bulletin. 

Idiocerus verticis (Say). 

(PI. XXV, fig. V.) 

Length, 1.55 mm.; greatest width, 0.15 mm. Apical portion only sHghtly 
wider than basal; slightly curv^ed, tip greatly narrowed with extreme apex 
rounded, chitinization light; strengthening rod extends caudad as far as 
fourteenth dorsal tooth. Toothed area on dorsal edge occupies a little more 
than the apical third ; teeth twenty in number, large, rounded, regular, evenly 
spaced except basally; bear no distinct secondary teeth, though the margin 
is irregularly roughened suggesting secondary' teeth; the tip bears ten small 
caudad-pointing teeth along its ventral edge; the ducts are inconspicuous, 
their circular openings alone being visible; of these apparently two open 
ventrally, three apicalty, and many dorsally; there is no distinct chitinized 
connection between the two valves of the pair. 

Idiocerus scurra (Germar) . 

(PI. XXV, fig. 8.) 

Length, 222 mm.; greatest width, 0.28 mm. About the same width for 
entire length; slightly cui-ved, tip nan-owed but broadly rounded at the ex- 
treme apex, rather heavily chitinized; strengthening rod extends caudad as 
far as the twenty-fifth, dorsal tooth. Toothed area on dorsal edge occupies 
a little more than the apical half; teeth thirty in number, moderately large, 
rounded, regular, evenly spaced, about the same distance apart for entire, 
length; bear no secondary teeth; the ventral edge of the tip bears ten small 
teeth; area of ducts conspicuous, granular in appearance, circular openings 
visible; two open ventrally, two apically, and many dorsally; the two valves 
of the pair are joined one to the other by a distinct, elongate, chitinized con- 
nection present on the dorsal edge of the basal area. 

Idiocerus ramentosus (Uhler) . 

(PI. XXV, fig. 4.) 

Length, 1.96 mm.; greatest width, 0.17 mm. About the same width for en- 
tire length; only slightly curved, tip greatly narrowed with extreme apex 
rounded, moderately heavy chitinization; strengthening rod extends caudad 
as far as eleventh dorsal tooth. Toothed area on dorsal edge occupies a little 
more than the apical third ; teeth eighteen in number, large, rounded, generally 
regular with a few double teeth, evenly spaced but farther apart basally; bear 
no secondary teeth; the tip bears three to four small, inconspicuous teeth 
along its ventral edge; area of ducts conspicuous, granular in appearance, 
circular duct openings visible; two to three open ventrally, three apically, 
and many doreally;, the two halves of the pair joined one to the other by 
an elongate, narrow, chitinized connection present on the dorsal edge of the 
basal area. 

Idiocerus snowi Gillette and Baker. 

(PI. XXV, fig. 3.) 

Length, 2.33 mm.; greatest width, 0.22 mm. About the same width for 
entire length; slightly curved, tip very bluntly rounded, rather heavily chitin- 
ized; strengthening rod extends caudad as far as eighth dorsal tooth. 
Toothed area on dorsal edge occupies the apical fourth; teeth ten in number, 



Readio: Ovipositors of Cicadellid.e. 227 

medium in. size, broad ami rounded, regularly spaced except basally, where 
they are farther apart; bear no secondary teeth; no teeth present at the 
apex, area of ducts conspicuous, granular in appearance, circular duct open- 
ings visible, none open ventrallj-, six apically, and many dorsally; the two 
valves of the pair are joined one to the other by an elongate, narrow, chitin- 
ized connection present on the doi-sal edge of the basal. area. 

Genus Macropsis Lewis. 
The ovipositors of two species of this genus have been examined 
and found to be very similar. They are long, narrow and rodlike, 
about the same width for entire length, slightly curved, and narrow 
only at the apex to an obtuse point. The two valves of the pair are 
not identical as to length, shape of tip, and teeth found at tip. The 
toothed area occupies only a small part of the length at the apex. 
The teeth are few in number, rather large, rounded, and may or 
may not bear secondary teeth. The tip is notched with small teeth 
in both species. The ducts are elongate, conspicuous, few in num- 
ber, and located only apically. 

Macropsis viridis (Fitch). 

(PI. XXVI, fig. 1.) 

Length, 2.3 mm.; greatest width, 0.05 mm. Very long, narrow, rodlike, 
about the same width for entire length, the two valves of the pair are not 
identical in length, shape of tip, and teeth found at tip; slighth' curved, nar- 
rows at apex to obtuse point, rather heavily chitinized, strengthening rod 
extends caudad as far as second dorsal tooth. Toothed area occupies but a 
small portion at the apex, the longer valve bears two large teeth on its dorsal 
edge and eleven small, inconspicuous teeth along its ventral edge, the shorter 
valve bears three large teeth along its dorsal edge and eleven small teeth 
along its ventral edge, none of these teeth bear secondaiy teeth; the extreme 
apex of the longer valve is entirely devoid of teeth, while the ventral teeth 
of the shorter valve extend to the apex, ducts are conspicuous, few in num- 
ber, elongate, all opening apically, two to four openings visible. 

Macropsis suturalis (Osborn and Ball). 

(PI. XXVI, fig. 2.) 

Length, 2.1 mm.; greatest width, 0.05 mm. Very long, narrow, rodlike, 
about the same width for entire length; the two valves of the pair are not 
identical, but differ in length, shape of tip, and teeth at tip; slightly curved, 
narrows at tip to an obtuse point, chitinization moderately heavy, some- 
what lighter than in M. viridis; strengthening rod extends caudad as far as 
fifth dorsal tooth. Toothed area includes only the apical portion ; the longer 
valve bears on its dorsal edge two large, heavily chitinized teeth at the tip 
and three large, somewhat less heavily chitinized teeth back from the tip; 
on its ventral edge are seventeen small teeth which are continuous around 
the tip; the shorter valve bears three large, heavily chitinized teeth at the 
tip, three less heavily chitinized teeth back from the tip, both groups being 



228 The University Science Bulletin. 

on the dorsal edge, and tliirteen small teeth on the ventral and apical edges; 
the three large teeth at the apex of the shorter valve bear small secondary 
teeth; the ducts are conspicuous, few in number, all located at the apex, six 
to seven openings visible. 

Oncopsis distinctus (Van Duzee). 

(PI. XXVI, fig. 3.) 

Length, 1.5 mm.; greatest width, 0.08 mm. Long, narrow, rodlike, about 
the same width for entire length; the two valves of the pair are not identical, 
but differ in length, shape of tip, and teeth found at tip; slightly curved, 
narrow at apex to obtuse point, heavily chitinized; strengthening rod ex- 
tends caudad as far as the second dorsal tooth. Toothed area on dorsal edge 
occupies only the apical portion of the valve; the longer valve bears two large, 
heavily chitinized teeth on the dorsal edge and fourteen small, inconspicuous 
teeth on the ventral edge, and the shorter valve bears two large, heavily 
chitinized teeth on the dorsal edge and fourteen small, inconspicuous teeth 
on its ventral and apical edges; none of the teeth bear secondary teeth; the 
ducts are conspicuous, elongate, few in number; five to six open apically, 
and others are scattered along the basal rod of the valve and open dorsally. 

Genus Bythoscopus Germar. 
The, ovipositors of two species of this genus have been examined 
and found to be simiUar. They are about the same width for the 
entire length; slightly though distinctly curved, narrow abrutly near 
the apex, and end in a rounded extreme apex. The teeth on the 
dorsal edge occupy about a third of the apical length, are numerous, 
small, irregular, and bear no secondary teeth. There are teeth on 
the ventral edge of tip in both species. The ducts are conspicuous, 
elongate, rather few in number. In both species there is present on 
the dorsal edge about midway of the valve an elongate, distinct, 
heavily chitinized connection between the two valves of the pair. 

Bythoscopus apicalis (Osborn and Ball). 

(PI. XXV, fig. 10.) 

Length, 1.53 mm.; greatest width, 0.14 mm. The apical portion is slightly 
wider than the basal portion, distinctly curved; the valve narrows from a 
point about six-sevenths of its entire length to the tip, extreme apex rounded, 
chitinization moderately heavy; strengthening rod extends caudad to within 
six dorsal teeth from the apex. Toothed area on the dorsal edge occupies a 
little more than the apical third of the length; teeth numerous, very small 
and inconspicuous, flatly rounded, irregular in shape and size, unevenly 
spaced; bear no secondary teeth; the ventral edge of the tip bears a few in- 
conspicuous teeth; ducts conspicuous, comparatively few in number, one group 
opens apically, and another group along the basal rod of the valve about mid- 
way; the two valves of the pair are joined one to the other by a distinct, 
elongate, chitinized connection present on the dorsal edge about midway of 
the valve, this connection bearing an angular tooth near its caudal extremity. 



Readio: Ovipositors of Cicadellid.e. 229 

Bythoscoptis miscellus (Stal). 

(PI. XXV, fig. 11.) 

Length, 1.27 iniu.; greatest width, 0.08 mm. About the same width for 
entire length; distinctlj^ curved, tip narrowed abruptly from a slight promi- 
nence on the dorsal edge about eight-ninths of the length to a broadly rounded 
e.xtreme apex; chitinization medium heavj', somewhat lighter than in B. 
apicnlis; strengthening rod extends caudad as far as fourteenth doi-sal tooth 
from the apex. Toothed area on dorsal edge occupies a little less than the 
apical third; teeth numerous, small, rather sharjily pointed, irregular, un- 
evenly si)aoed: bear no secondary' teeth; the tip is notched with teeth which 
are continuous around the tip, ten to eleven on the ventral edge; ducts con- 
spicuous, elongate, few in number; one group opens apically, another at the 
basal end of the toothed area, and another in the basal rod about midway; 
the tw^o valves of the pair are joined one to the other by a distinct, elongate, 
chitinized connection jiresent on the dorsal edge about midway. 

Subfamily CICADELLINiE Van Duzee. 

Genus Oncometopia Stal. ^ 

The ovipositors of two species of this genus have been examined 
and found to be simihir, though having some very distinct differ- 
ences. The ovipositor of 0. iindata more closely resembles the 
ovipositor of Homalodisca triquetra than it does the ovipositor of 
0. lateralis. In the ovipositors of this genus we find a narrow, 
curved, rodlike basal attachment, and a broad, fiat, apical shaft 
bearing teeth along its dorsal edge, and a preapical prominence on 
its ventral edge. The primary teeth in each case bear secondary 
teeth. The tip is notched with small teeth on both edges; the ducts 
are conspicuous, elongate, numerous, and distinctly curved. 

Oncometopia' undat a (Fabricius) . 

(PI. XXII, fig. 3; pi. XXVI, fig. 4.) 

Length of toothed area, L8 mm.; greatest width, 0.27 mm. Narrow and 
rodlike at base, widens beyond base into flat, broad apical shaft, which is 
about the same width for entire length and tapers only at the apex; pre- 
apical prominence present, conspicuous, broadly rounded; apical shaft slightly 
curved, tip broadly rounded, chitinization medium; strengthening rod extends 
caudad as far as last doi-sal tooth. Toothed area on dorsal edge occupies 
entire length of broadened shaft; teeth thirty-one to thirty-two in number, 
of medium size, of a general triangular shape with the caudal side much 
longer than the cephalic side; apices rounded, regular, evenly spaced, more 
heavily chitinized than the rest of the valve; possess secondary teeth on both 
caudal and cephalic edges, one to four on cephahc edge and two to five on 
caudal edge, those on the caudal edge much larger than those on the cephalic 
edge; tip notched with small teeth on both dorsal and ventral edges, not con- 
tinuous around the tip, fifteen teeth present on ventral edge between preapical 



230 The University Science Bulletin. 

prominence and extreme apex; ducts conspicuous, decidedly cui-ved, numerous, 
most numerous apically; open apically and along dorsal edge in and near 
teeth, one to three to each tooth. 

Oncometopia lateralis (Fabricius). 

(PI. XXII, fig. 4; pi. XXVI, fig. 5.) 

Length 1.66 mm.; greatest width, 0.2 mm. Narrow and rodlike at the base, 
widens beyond base into broad, flat apical shaft, which tapers very slightly 
toward apex; preapical prominence present, rather indistinct, broadly rounded; 
only very slightly cui-ved beyond the base, tip broadly rounded, chitinization 
medium, strengthening rod extends caudad as far as last dorsal tooth. Toothed 
area on dorsal edge occupies the entire length of the apical shaft; teeth eigh- 
teen to twenty in number, large, subquadrate in shape with the cephalic side 
longer than the caudal and bearing a smoothly rounded prominence, regular, 
evenly spaced, more heavily chitinized than the rest of the valve and have 
the appearance of being set into the rest of the valve; bear secondary teeth, 
seven to fifteen on the broad outer edge and four to eight on the cephalic 
side, those on the outer edge larger than those on the cephalic side; the tip 
is notched with small teeth on both dorsal and ventral edges, not continuous 
around the tip, twenty-one teeth between preapical prominence and extreme 
apex; ducts conspicuous, elongate, distinctly curved, numerous, most numerous 
apically ; open apically, along the dorsal edge in and near teeth, two to four to 
each tooth; first three teeth not served with ducts. 

Homalodisca triquetra (Fabricus). 

(PI. XXII, fig. 5; pi. XXVI, fig. 6.) 

Length, 2.6 mm.; greatest width, 0.33 mm. Narrow and rodlike at base, 
widens beyond base into broad, flat portion, which is about the same width 
for entire length, narrowing only at apex; preapical prominence present, 
distinct, broadly rounded; uncurved beyond base, extreme apex broadly 
rounded, chitinization medium; strengthening rod extends caudad as far as 
third dorsal tooth from the apex. Toothed area on dorsal edge occupies the 
entire length of the apical shaft; teeth forty-four to forty-five in number, 
medium in size, of a general triangular shape with the caudal side longer than 
the cephalic, regular in size and shape, evenly spaced, more heavily chitinized 
than the rest of the valve ; bear secondary teeth, three to seven on the cephalic 
edge and five to nine on the caudal edge; those on the caudal edge are larger 
than those on the cephaUc edge; tip notched with small teeth on both dorsal 
and ventral edges, not continuous around the tip, ducts conspicuous, numerous, 
slightly curved, most numerous apically; open apically along the dorsal edge 
in and near teeth, two to five to each tooth; the first six teeth not served by 
ducts. 

Aulacizes irrorata (Fabricius), 

(PI. XXII, fig. 6; pi. XXVI, fig. 7.) 

Length, 2.9 mm.; greatest width, 0.44 mm. Narrow and rodlike at the base, 
widens beyond base into broad, flat, apical shaft; reaches point of greatest 
width about two-fifths of length and from that point tapers gradually to the 
tip; preapical prominence present, conspicuous, obtuse-angled; not curved 



Readio: Ovipositors of Cicadellid.e. 231 

be5'ond curved basal attachmont; tip bluntly rounded, chitinization medium; 
strengthening rod extends caudad as far as third dorsal tooth from apex. 
Toothed area on dorsal edge occupies the entire length of the apical shaft; 
teeth forty-one to fortj'-thrce in number, rather small, in the shape of a flat- 
tened isosceles triangle, regular in size and shape, evenly spaced; the first 
twelve teeth bear no secondary teeth; the teeth caudad of the first twelve teeth 
bear four to ten secondary teeth on the caudal side; tip notched with small 
teeth on both dorsal and ventral edges, not continuous aroimd the tip; thirty- 
five to thirtj'-seven teeth on ventral edge between preapical prominence and 
extreme apex; ducts conspicuous, elongate, rather narrow, curving, numerous, 
most numerous apically; open apically, along dorsal edge, near but not in 
teeth, and in face of valve back from edge; the first twelve teeth are not 
served by ducts. 

Genus Cicadella Latreille. 

The ovipcsitors of two species of this genus have been examined 
and found to be similar. In each case the ovipositor consists of a 
narrow, curved basal attachment and a wider apical shaft which 
bears teeth along its dorsal edge, in turn bearing secondary teeth. 
A preapical prominence is present in one species, absent in the other. 
The tip is notched on both dorsal and ventral edges. The ducts 
are rather inconspicuous, elongate, rather few in number, and dis- 
tinctly curved. 

Cicadella hicroglyphica (Say). 

(PI. XXII, fig. 7; pi. XXVI, fig. 9.) 

Length, 1.66 mm.; greatest width, 0.13 mm. Curved basal attachment nar- 
row and rodlike; apical portion consists of a broad, flat shaft, about the same 
width for entire length, tapering only at apex, preapical prominence present 
on the ventral edge, obtuse-angled; onl}- slightly curved; apex greatly nar- 
rowed and acutely pointed, very lightly chitinized; strengthening rod ex- 
tends caudad as far as last dorsal tooth. Toothed area on dorsal edge occu- 
pies the entire length of the apical shaft ; teeth seventeen to eighteen in num- 
ber, medium in size, somewhat triangular with the apex flattened and the 
caudal side slightly longer than the cephalic, regular in size and shape, evenly 
spaced, more heavily chitinized than the rest of the valve; bear a large num- 
ber of small, irregular, secondary teeth on both edges; tip notched with small 
teeth on both edges; a large number of very small teeth occur between the 
tip of the preapical prominence and the extreme apex; ducts rather incon- 
spicuous, few in number greatly curved; open apically, along dorsal edge in 
and near teeth, and a few in the face of the valve back from the edge. 

Cicadella circellata (Baker). 

(PI. XXII, fig. 8; pi. XXVI, fig 8.) 

Length, 1.6 mm.; greatest width, 0.09 mm. Apical, flattened portion only 
slightly wider than basal portion; preapical prominence wanting; only slightly 
curved, tip greatly narrowed but rounded at extreme apex, chitinization 
rather light but heavier than in C. hieroglyphica; strengthening rod extends 
caudad as far as and beyond last dorsal tooth. Toothed area on dorsal edge 



232 The University Science Bulletin. 

occupies nearly all of broadened area or about half the total length; teeth 
thirteen in number, rather small, in the general shape of a greatly flattened 
obtuse triangle with the caudal side much longer than the cephalic, regular 
in spacing; bear secondary teeth on both edges, five to eight on caudal edge 
and one to two on cephalic edge, those on the caudal edge are much the 
larger; tip notched with small teeth on both dorsal and ventral edges, not con- 
tinuous around the tip, twenty present on ventral edge; ducts rather incon- 
spicuous, though more conspicuous than in C. hieroglyphica, elongate, few in 
number, distinctly curved; open apically, along dorsal edge in and near teeth, 
and in face of valve back from edge. 

Genus Kolla Distant. 

The ovipositors of three species of this genus have been examined 
and found to be similar. In each case the ovipositor consists of a 
curved, narrow, rodhke basal attachment and a broader, flat, apical 
shaft bearing teeth along its dorsal edge. The teeth are triangular 
in shape and bear small secondary teeth along their caudal sides. 
The tip is notched in each case with small teeth present on both 
edges, but not continuous around the tip. The ducts may or may 
not be visible; when visible they are elongate, rather straight, and 
few in number. 

Kolla bifida (Say). 

(PI. XXII, fig. 13; pi. XXVIII, fig. 3.) 

Length, 1.5 mm.; greatest width, 0.22 mm. Basal connection narrow and 
rodlike ; apical shaft broad, flat, narrowing gradually to the apex ; no preapical 
prominence ; only very slightly curved beyond curved basal attachment ; apex 
narrowed with extreme apex rounded, only lightly chitinized; strengthening 
rod extends caudad as far as and beyond last apical tooth. Toothed area on 
dorsal edge occupies entire length of apical shaft and more than two-thirds of 
the entire length; teeth fourteen in number, medium in size, in the general 
shape of a greatly flattened obtuse triangle with the caudal edge much longer 
than the cephalic and the apex rounded, not distinctly more heavily chitinized 
than the rest of the valve; bear small secondary teeth along the caudal edge 
only, seven to twenty-one on each tooth; the tip is notched with small teeth 
on both dorsal and ventral edges, not continuous around the tip, fourteen to 
fifteen on ventral edge; ducts inconspicuous, only apices and circular open- 
ings visible; open apically, along the doi-sal edge in and near teeth, and a few 
in the face of the vahe back from the edge. 

Kolla geometrica (Signoret). 

(PI. XXVIII, fig. 2.) 

Length, 1.17 mm.; greatest width, 0.17 mm. Basal portion narrow and rod- 
like; apical shaft broad, flat, tapers to apex; no preapical prominence; not 
curved bej'ond base; tip greatly narrowed with extreme apex rounded bluntly; 
chitinization light though somewhat heavier than in K. bifida; strengthening 
rod extends caudad as far as last dorsal tooth. Toothed area on dorsal 
edge occupies entire length of broadened shaft; teeth seventeen to nineteen in 



Readio: Ovipositors of Cicadellid.e. 233 

number, rather small, in the general shape of a greatly flattened obtuse tri- 
angle with the caudal side much longer than the cephalic and the apex rounded, 
rather irregular in size and spacing, slightly more heavily chitinized than the 
rest of the valve; bear secondary teeth along the caudal edge only, four to 
fourteen on each tooth; tip notched with teeth on both dorsal and ventral 
edges, not continuous around the tip, thirteen to fifteen on ventral edge; ducts 
easily visible for entire length, comparatively few in number, straight, more 
numerous apically; open apically, along dorsal edge in and near teeth, and in 
the face of valve back from edge. 

• Kolla hartii (Ball). 

(PI. XXII, fig. 14; pi. XXVIII, fig. 1.) 

Length, 1.4 mm.; greatest width, 0.22 mm. Basal portion narrow and rod- 
like; apical shaft broad and flat, tapers towards apex; no preapical promi- 
nence; not curxed beyond base, tip greatly narrowed, extreme apex smoothly 
rounded, lightly chitinized; strengthening rod extends caudad beyond last 
doi-sal tooth. Toothed area on dorsal edge occupies entire length of apical 
shaft; teeth seventeen in number, medium in size, in the shape of an obtuse 
triangle, though not so flat as in A', bifida and A', geometrica; caudal edge 
somewhat longer than cephalic edge, rather irregular in size and spacing, not 
more hea\'iiv chitinized than the rest of the valve; possess small secondary 
teeth on the caudal edge of each primaiy tooth, eight to seventeen to each 
tooth; tip notched with small teeth on both dorsal and ventral edges, not con- 
tinuous around tip, twenty-three on ventral edge; ducts more conspicuous than 
in K. bifida, less conspicuous than in A. geometrica; bases and apices visible 
but middle portions invisible; open apicalh', along dorsal edge in and near 
teeth, and in face of valve back from edge. 

Helochara communis Fitch. 

(PI. XXII, fig. 10; pi. XXVII, fig. 5.) 

Length. L44 mm.; greatest width, 0.22 mm. Basal portion narrow and 
ciu-ved, apical shaft broad and flat, about the same width for entire length, 
tapers toward tip; bears no preapical prominence; not curved beyond the 
base, tip greatlj^ narrowed, with the extreme apex ending in an obtuse point, 
chitinization mechimi; strengthening rod extends caudad to a point midway 
between the last dorsal tooth and the tip. Toothed area on dorsal edge oc- 
cupies the entire length of the apical shaft, nearly the entire length of the 
valve; teeth thirty in number, rather large, the first fourteen of a general 
triangular shape, apical sixteen subquadrate with the cephalic side longer than 
the caudal, rather irregular in size, shape, and spacing; basal fourteen primary 
teeth bear small secondaiy teeth on the two exposed sides of the triangle; 
apical sixteen primary teeth also bear secondary teeth on the outer margin, 
three to fourteen in number; tip is notched with small teeth on both doi^sal 
and ventral edges, not continuous around the tip, fort5'-six present on venti-al 
edge; ducts conspicuoiLS, rather numerous, elongate, rather straight, occupy 
entire shaft; open apically, along dorsal edge in and near teeth, and in face 
of valve back from edge. 



234 The University Science Bulletin. 

■4 

Graphocephala coccinca (Forster). 

(PI. XXII, fig. 9; pi. XXVIII, fig. 4.) 

Length, 1.87 mm.; greatest width, 0.22 mm. Basal portion narrow and rod- 
hke, apical shaft broad and flat, about the same width for entire length; pre- 
apical prominence present, distinct, obtuse-angled; not curved bej-ond base, 
tip greatlj' narrowed and ending in a rather blunt point, chitinization light; 
strengthening rod e.xtends caudad beyond last dorsal tooth. Toothed area on 
dorsal edge occupies entire length of apical shaft; teeth nineteen in number, 
medium in size, of a general triangular shape with the caudal side longer than 
the cephahc, regular in size and shape, distinctly more heavily chitinized than 
the rest of the valve; bear small secondarj^ teeth on both edges, three to six 
on cephalic edge and eight to seventeen on caudal edge; tip notched with 
small teeth on both dorsal and ventral edges, not continuous around the tip, 
forty-five between preapical prominence and extreme apex; ducts easily vis- 
ible though not conspicuous, elongate, rather few in number, curved, most 
numerous apically; open apically, along dorsal edge in teeth, one to two to 
each tooth, and a few in the face of valve back from edge. 

Genus Dr^culacephala Ball. 

The ovipositors of three species of this genus have been examined 
and found to be similar. The ovipositor of Helochara communis 
also shows its close relationship to this genus. In each case the ovi- 
positor consists of a curved, rodlike basal attachment and a broad, 
fiat, apical shaft which bears teeth along its dorsal edge for the en- 
tire length. The distal teeth are triangular and the apical teeth sub- 
quadrate. Both types of teeth bear small secondary teeth. The tip 
in every case is notched with small teeth on both dorsal and ventral 
edges, not continuous around the tip. The ducts may or may not 
be conspicuous, but where visible are elongate, numerous and curved. 

Drceculacephala mollipes (Say). 

(PI. XXII, fig. 13 ; pi. XXVII, fig. 2.) 

Length, 2.1 mm.; greatest width, 0.33 mm. Basal portion narrow and rod- 
like, apical shaft broad and flat, about the same width for entire length, nar- 
rowing caudad to apex; preapical prominence wanting; not curved beyond 
base, tip greatly narrowed by curving ventral edge and ends in an obtuse point, 
chitinization moderately heavy; strengthening rod extends caudad to a point 
midway between last dorsal tooth and apex. Toothed area on dorsal edge 
occupies entire length on apical shaft; teeth twenty-eight in number, large, 
basal nine of a general triangular shape, apical nineteen subquadrate with 
the cephalic side longer than the caudal, rather irregular in size, shape and 
spacing; the basal nine bear small secondary teeth on both cephalic and 
caudal edges; the apical subquadrate teeth also bear secondary teeth, seven 
to seventeen on the outer edge and all except the apical five bear secondary 
teeth along the cephalic edge, one to six in number; tip is notched with small 
teeth on both dorsal and ventral edges, not continuous around the tip, forty- 



Rkadio: Ovipositors of Cicadellid.^. 235 

two on the vent ml cnlge; ducts consijicuous, rather numerous, elongate, slightly 
curved, most numerous apicalh-; open apically, along dorsal edge in and near 
teeth, and in the .•^in-face of the valve back from edge. 

Drceculaccyh a la n o ve boracensu ( Fitch ) . 

(PI. XXVII, fig. 1.) 

Length, 2.65 mm.; greatest width, 0.33 mm. Narrow and rodlike at base, 
broad and flat ajiically, apical shaft about the .same width for entire length; 
prcapical prominence wanting; only slightly curved beyond base, tip greatly 
narrowed by slightly curving dorsal edge and greatly curving ventral edge 
and ends in a rather sharp point, chitinization light, much lighter than in the 
other members of this genus examined; strengthening rod extends cavidad to 
a point midwaj' between last dorsal tooth and apex. Toothed area on dorsal 
edge occupies entire length of apical shaft, the dorsal edge bearing the teeth 
more heavilj^ chitinized than the rest of the valve; teeth thirty-nine to forty- 
one in number, of medium size, the basal sixteen of a general triangular shape, 
the apical twenty-three subquadrate wath the cephalic side longer than the 
caudal, rather irregular in size, shape, and spacing; the basal triangular teeth 
bear small secondary teeth on both dorsal and ventral edges; the apical sub- 
quadrate teeth bear three to twelve secondary teeth on the outer edge and 
two to five on the cephalic edge; the secondary teeth midway of the valve 
are very small and indistinct; tip notched with small teeth on both dorsal 
and ventral edges, not continuous around the tip, forty-five present on the 
ventral edge; ducts inconspicuous, very faintly visible except at openings, 
rather numerous, most numerous apically; open apically, along dorsal edge 
in and near teeth, and in surface of valve back from edge. 

Drceculacephala reticulata (Signoret). 

(PI. XXII, fig. 12; pi. XXVII, fig. 3.) 

Length, L75 mm.; greatest width, 0.27 mm. Basal portion narrow and rod- 
like, apical portion broad, fiat, about the same width for entire length, nar- 
rows at apex; preapical prominence wanting; not curved beyond curved basal 
attachment, tip greatly narrowed by slightly .curved dorsal edge and greatly 
curved ventral edge, ends in obtuse point, chitinization medium, less heavy 
than in D. noveboraccnsis and heavier than in D. mollipes; strengthening rod 
extends caudad to a point midwaj^ between last dorsal tooth and apex. 
Toothed area on dorsal edge occupies apical shaft for entire length; teeth 
twenty-six to twenty-eight in number, rather large, basal eleven of a general 
triangular shape, apical fifteen subquadrate with the cephalic side longer 
than the caudal, rather irregular in size, shape and spacing; basal eleven bear 
small secondai-y teeth on both edges; apical subquadrate teeth also bear sec- 
ondary teeth, three to eleven on the outer edge and, with the exception of 
the apical three, one to five on the cephalic edge; tip notched with small 
teeth on both dor.sal and ventral edges, not continuous around the tip, 42 pres- 
ent on the ventral edge; ducts conspicuous, rather straight, numerous, most 
numerous apically; open apically, along dorsal edge in and near teeth, and a 
few in the face of valve back from edge. 



236 The University Science Bulletin. 

Pagaronia tripunctata (Fitch). 

(PL XXVII, fig. 6.) 

Length, 1.5 mm.; greatest width. 0.17 mm. Narrow and rodhke at base, 
apical shaft broad and flat, about the same width for entire length, narrows 
caudad to aj^ex; preapical prominence wanting; not curved beyond base, tip 
narrowed, bkmtly rounded at extreme apex, chitinization Hght ; strengthening 
rod extends caudad as far as last dorsal tooth. Toothed area on dorsal edge 
occupies entire length of apical shaft, dorsal edge bearing teeth is more heavily 
chitinized than the rest of the valve; teeth seventeen to twenty in number, 
medium in size, in the shape of a flattened obtuse triangle with the caudal 
side much longer than the cephalic, irregular in size, shape and spacing; bear 
secondary teeth on both outer edges, three to ten on cephalic edge and three to 
twenty-five on the caudal edge; tip notched with small teeth on both doreal 
and ventral edges, not continuous around the tip, 30 present on ventral edge; 
ducts conspicuous, rather numerous, only slightly curved, most numerous 
apically; open apically, along dorsal edge in and near teeth, and a few in the 
surface of valve back from edge. 

Errhomenellu.s montanus Baker. 

(PI. XXVII, fig. 4.) 

Length, 2.83 mm.; greatest width, 0.22 mm. About the same width for en- 
tire length; preapical prominence present, rather indistinct, broadly rounded; 
only slightly curved beyond curved basal attachment, tip greatly narrowed 
with the extreme apex rounded, chitinization medium; strengthening rod ex- 
tends caudad as far as fourth dorsal tooth. Toothed area on dorsal edge oc- 
cupies less than a fourth of the entire length apically; teeth seven to eight in 
number, small, inconspicuous, rounded, regular in size and shape, rather far 
apart, uneven in spacing, bear no secondary teeth; tip bears no teeth; no dis- 
tinct ducts visible, circular openings plainly visible, of these a number occur 
on the ventral apical edge, some at the extreme apex, and also along the en- 
tire dorsal edge. 

Subfamily GYPONINtE (Stal). 
Genus Gypona Germar. 

The ovipositors of four species of this genus have been examined 
and found to be similar. The ovipositor of Xerophloea liridis, the 
only other species of this subfamily examined, is not similar to the 
ovipositors of the genus Gypona. In this genus the ovipositor is 
relatively short, stout and broad, usually being rather heavily chi- 
tinized. The apical portion is not much, if any, wider than the basal 
portion. Teeth are present along the dorsal edge for from one- 
fourth to one-half the apical length. The tip may or may not bear 
teeth on one or both edges. The ducts in every case are conspicuous, 
large, elongate and numerous. 



Readio: Ovipositors of Cicadellid.^. 237 

Gypona octo-lineata (Say). 

(PI. XXIII, fig. 1; pi. XXVIII, fig. 8.) 

Length, 2 iiun.; greatest width, 0.48 mm. Consists of a rather narrow basal 
portion, which widens out into a broad, apical portion, which narrows caudad 
to the apex, point of greatest width about midway; no preapical prominence; 
shghtiy curved, tip greatly narrowed with the extreme apex broadly rounded, 
chitinization moderately heavy; strengthening rod extends caudad almast to 
apex. Toothed area on dorsal edge occupies about a half of the apical length; 
teeth numerous, rather small, rounded, irregular in size, shape- and spacing; 
bear no secondary teeth; tip bears small teeth on both dorsal and ventral 
edges, not continuous around the tip, thirteen on ventral edge; ducts con- 
spicuous, large, numerous, apparently arising from a common cavity in the 
center of the valve, most numerous at the tip; open along ventral apical edge, 
at extreme apex, and along the dorsal edge for the entire length, though very 
few in basal region. 

Gypotia biniaculata Spangberg. 

(PI. XXVIII, fig. 5.) 

Length, 3.17 mm.; greatest width, 0.42 mm. About the same width for en- 
tire length, bears a broadly rounded, heavily chitinized prominence on the dor- 
sal edge somewhat less than midway of the length, tapers caudad toward apex; 
slightly curved, tip gradually naiTOwed, ending in a blunt point, chitinization 
hea\-y, much hea\ ier than in G. octo-lineata; strengthening rod extends caudad 
almost to apex. Toothed area on dorsal edge occupies about one-third of 
apical length; teeth numerous, rather small, rounding, iiTegular in size and 
shai)e; bear no secondaiy teeth; tip notched on dorsal edge and at extreme 
apex with small teeth, none present on ventral edge; ducts conspicuous, very 
large, rather straight; open apically and along dorsal and ventral edges for 
the length of the toothed area, a few also open in the basal region. 

Gypona angulata Spangberg. 

(PI. XXVIII, fig. 6.) 

Length, L75 mm.; greatest width, 0.35 mm. Apical portion somewhat wider 
than basal, point of greatest width about midway, tapers caudad to apex; 
only slightly curved, tip gradually narrowed with the extreme apex rounded, 
chitinization moderately heavy; strengthening rod extends caudad to within 
a short distance of tip. Toothed area extends from the prominence on the 
dorsal edge to the apex; teeth numerous, small and indistinct, rather flatly 
rounded, verj' irregular in size, shape and spacing; bear no secondary teeth; 
no teeth on tip; ducts consi;Hcuous, large, numerous, broadly curving, most 
numerous apicall}^; open apically, along ventral apical edge, along dorsal 
edge of toothed area, and a few in the basal portion of the valve. 

Gypona pectoralis Spangberg. 

(PI. XXVIII, fig. 7.) 

, Length, L83 mm.; greatest width, 0.42 mm. Very broad and heavy for 
entire length, point of greatest width about midway, where there is a broadly 
rounded prominence on the dorsal edge, narrows caudad to apex; slightly 
curved, tip greatly narrowed with extreme apex rounded, chitinization very 



238 The University Science Bulletin. 

heavy; strengthening rod extends caudad almost to apex. Toothed area on 
dorsal edge occupies about a fourth, of the apical length ; teeth very few in 
number, small, flat, somewhat rounded, very irregular in size, shape and 
spacing, bear no secondary teeth; no teeth present at extreme apex; ducts 
conspicuous, large, somewhat branching, more numerous apically; open 
apically, along ventral apical edge, and along dorsal for entire length. 

Xerophloea viridis (Fabricius). 

(PL XXIII, fig. 2; PI. XXIX, fig. 2.) 

Length, 2.88 mm.; greatest width, 0.08 mm. Narrow and rodlike, about the 
same width for entire length; distinctly curved, tip bluntly rounded, chitiniza- 
tion medium; strengthening rod extends caudad as far as twentieth dorsal tooth. 
Toothed area on dorsal edge occupies a little less than the apical third; teeth 
twenty-four in number, medium in size, rounded, regular in size and shape, 
the apical four are farther apart than the basal twenty; the apical three bear 
one to three secondary teeth on the caudal edge; tip notched with small teeth 
on both dorsal and ventral edge, not continuous around the tip, twelve to 
thirteen on ventral edge; ducts conspicuous, relatively few in number, most 
numerous at apex; open apically and along entire dorsal edge, even in basal 
portion. 

Subfamily JASSIN./E (Amyot and Serville). 

Tribe Acucephalini (Dohrn). 

Stroggylocephalus agrestis (Fallen). 

(PI. XXIX, fig. 3.) 

Length, L88 mm.; greatest width, 0.17 mm. Curved base rather narrow 
and rodlike, rather broad and flat beyond base, about the same width for 
entire length, tapers gradually toward apex; curved at base of broadened area, 
tip greatly narrowed with extreme apex narrowly rounded, chitinization rather 
light, strengthening rod extends caudad as far as sixth dorsal tooth from apex. 
Toothed area on dorsal edge occupies a little less than half the entire length, 
teeth twenty-six to twenty-eight in number, rather small, of a general tri- 
angular shape, some roimded and some sharply pointed, verj' irregular in size, 
shape and spacing; bear no secondary teeth; tip notched with a few small, 
indistinct, irregular teeth on the ventral edge only; ducts conspicuous, elongate ; 
open along ventral apical edge, at extreme apex, and along entire dorsal edge; 
the two valves of the pair are joined one to the other by a distinct, elongate, 
heavily chitinized connection present on the dorsal edge near the base of the 
broadened area. 

Memnonia consobrina Ball. 

(PI. XXIX, fig. 5.) 

Length, 2.2 mm.; greatest width, 0.25 mm. Gunned basal attachment nar- 
row and rodlike, beyond base is a rather narrow, lightly chitinized area ex- 
tending caudad for about a third of the length; caudad of this is a broader, 
more heavily chitinized portion occupying the rest of the valve; narrows, 
rather abruptly caudad to apex; only slightly curved beyond base, tip greatly 
narrowed by broadly cui-ving doi-sal edge; extreme apex ends in sharp, acute- 



Readio: Ovipositors of Cicadellid.e. 239 

angled point, rhitiuization nioderatt'ly light; strengthening rod extends caudad 
almost to apex. There are no evidences of distinct teeth, though the margin 
is irregularly roughened in se\eral i)laces; area of ducts conspicuous, granular 
in appearance, circular duct ojienings visible; ope>n along ventral apical edge, 
at extreme apex, and along dorsal edge for entire length. 

Xestocephahis pnlicarius Van Duzee. 

(PI. XXIII, fig. 3; pi. XXIX, fig. 1.) 

Length. 0.88 mm.; greatest width, 0.08 mm. Curved basal attachment 
narrow and rodlike, rest of valve about the same width for entire length; 
preapical prominence on ventral edge present, distinct, obtuse angled, giv- 
ing the apical portion of the valve the appearance of a spear head; tip 
narrowed abruptly beyond preapical prominence; very distinctly, broadly 
and evenly curved; tip greatly narrowed, ending in sharply pointed ex- 
treme apex, chitinization medium ; strengthening rod extends caudad to 
within a short distance of extreme apex; dorsal edge of tip bears an indistinct 
notch where rod meets dorsal edge. Toothed area on dorsal edge occupies 
the apical three-fourtlxs of the length; teeth fourteen to fifteen in number, 
small, indistinct, rather irregular in size, shape and spacing, especially basally; 
bear no secondary teeth; tip bears no teeth; ducts rather inconspicuous though 
plainly visible, rather few; open in preapical prominence, at extreme apex, 
and along entire dorsal edge; the two valves of the pair are joined one to 
the other by a distinct, elongate, narrow, heavily chitinized connection present 
on the dorsal edge, this connection more liberally supplied with ducts than the 
portion of the valve immediately caudad of it. 

Tribe Jassini (Dohrn). 
Dorycephalus platyrhynchus Osborn. 

(PI. XXIX, fig. 4.) 

Length, 3.5 mm.; greatest width, 0.5 mm. Curved basal portion narrow 
and rodlike ; at a point about a third of the length it widens into a broad, 
flat shaft which tajiers slightly caudad; point of greatest width is a little less 
then midway; only slightly cui-ved beyond base, tip greatly narrowed by 
rounding dorsal edge, extreme apex rounded, chitinization very light; strength- 
ening rod extends caudad to within a short distance of the extreme apex. 
There are no distinct teeth, though the ventral apical edge and the entire 
dorsal edge are slightly and irregularly roughened; area of ducts inconspicu- 
ous, no ducts visible, circular duct openings visible; a single row opens along 
the ventral apical edge, a single row along the dorsal apical edge, distad 
of these a double row, distad of the double row a triple row, and still further 
distad in the wider portion of the valve as many as four or five irregular rows 
open in the entire breadth of the valve. 

Hecalus lineatus (Uhler). 

(PI. XXIX, fig. 8.) 

Lenth, 4.1 mm.; greatest width, 0.43 mm. Curved basal portion narrow and 
rodlike, widens bej'ond base into broa<l, flat portion which tapers gradually 
caudad to apex, point of greatest width about two-fifths of length; only slightly 
curved bej'ond base, tip greatly narrowed, ending in sharp point chitinization 



240 The University Science Bulletin, 

moderately light; strengthening rod extends caudad to within a short distance 
of the apex. No distinct teeth present, the ventral apical edge and the entii-e 
dorsal edge except at the apex are slightly and irregularly roughened; area of 
ducts conspicuous, granular in appearance, circular duct openings visible; open 
along ventral apical edge and along dorsal edge for entire length, in the basal 
region they open irregularly in the entire width of the valve. 

Sipanghergiella mexicana Baker. 

(PI. XXIX, fig. 6.) 

Length, 1.9 mm., greatest width, 0.27 mm. Cvu-ved base narrow and rodlike, 
widens into broad, flat portion, about the same width for entire length, nar- 
rowing caudad to apex; bears two prominences, one on dorsal edge somewhat 
less than midway, and the other on the ventral edge about two-thirds the 
length, more angular; not curved beyond the base, narrowed at tip evenly and 
gradually by both curving edges, ends in sharp point, chitinization medium; 
strengthening rod extends caudad almost to apex. Bears no teeth; area of 
ducts granular in appearance, circular openings very small and inconspicuous; 
open along ventral apical edge, at extreme apex, along dorsal edge, and in basal 
portion of valve. 

Parabolocratus ^avidus Signoret. 

(PI. XXIX, fig. 7.) 

Length, 2.3 mm.; greatest width, 027 mm. Curved base narrow and rodlike, 
widens gradually into broad, flat portion, about the same width for entire 
length, narrowing only at apex; bears two broadly rounded prominences, one 
on the dorsal edge about a third of the length, the other on the ventral edge 
about two-thirds the length; only slightly curved beyond base, tip gradually 
narrowed and ends in sharp point, chitinization medium; strengthening rod 
extends caudad to within a short distance of the apex. Toothed only at ex- 
treme tip; teeth are continuous around the tip, of these six are on the dorsal 
edge and ten on the ventral edge; ducts inconspicuous, invisible except at ex- 
treme apices, circular openings visible ; open along ventral apical edge and 
along entire dorsal edge except in dorsal prominence. 

Aligia jucunda (Uhler). 

(PI. XXIX, fig. 11.) 

Length, L83 mm.; greatest width, 0.17 mm. Curved base narrow and rod- 
like, slightly wider beyond base for a little more than half the length; apical 
portion of valve widens abruptly, tapers caudad to apex; distinctly curved, tip 
narrowed by curved dorsal edge, extreme apex bluntly rounded, almost square, 
chitinization medium; strengthening rod extends caudad to within fourteen 
dorsal teeth of apex. Toothed area on dorsal edge occuj^ies somewhat less than 
the apical half; teeth eighty-eight in number, very small, wedge-shaped, fairly 
regular in size, shape and spacing, bear small secondaiy teeth on outer edge, 
one to four in number; tip devoid of teeth on ventral and caudal edges; area 
of ducts conspicuous, granular in appearance, though a few elongate ducts are 
visible apically; ducts straight, numerous; open along ventral apical edge, at 
extreme apex, along dorsal edge of toothed area, and irregularly in the basal 
region. The two valves of the pair are joined one to the other by an elongate 



Readio: Ovipositors of Cicadellid.e. 241 

connection only slightly more heavily chitinized than the rest of the valve, 
present on the dorsal edge in the basal region. 

Genus Mesamia Ball. 

Two species of this genus have been examined and found to be 
similar. In each case the ovipositor consists of a curved basal at- 
tachment, a somewhat wider portion that extends about to the mid- 
point, and an apical portion which bears teeth along its dorsal edge. 
A small preapical prominence is present on one of the two species. 
Some of the primary teeth, at least, bear secondary teeth. In one 
species the tip bears distinct teeth, in the other the tip is only irreg- 
ularly roughened. The area of ducts is granular in appearance and 
the duct openings are visible. The two valves of the pair are joined 
one to the other by an elongate, heavily chitinized connection pres- 
ent on the dorsal edge in the basal region. 

Mesamia straminea (Osborn). 

(PI. XXIX, fig. 9.) 

Length, 1.53 mm.; greatest width, 0.18 mm. Curved basal portion narrow 
and rodlike, bej^ond base is a wider, heavity chitinized area extending almost to 
the midpoint; the apical toothed area occupies the rest of the valve and is 
slightly wider, less heavily chitinized, and tapers caudad to the apex; bears no 
preapical prominence; slightly, though distinctly curved, tip narrowly rounded, 
chitinization moderately heavy; strengthening rod extends caudad as far as 
fourth dorsal tooth from the tip. Toothed area on dorsal edge occupies the 
apical half; about twenty-five primaiy teeth present, of medium size, very 
irregular as to size, shape and spacing; the apical ten are somewhat regular, 
long and flatly rounded and may or ma}^ not bear a few secondaiy teeth on the 
caudal edge, the teeth distad of these extremely irregular; the tip is irregularly 
roughened on both dorsal and ventral edges, but bears no distinct teeth; area 
of ducts granular in appearance, duct openings easily visible; open along ven- 
tral apical edge, at extreme apex, along dorsal edge of toothed area, and in the 
basal portion of the valve; the two valves of the pair are joined one to the 
other by a distinct, elongate, heavily chitinized connection present on the 
dorsal edge at about one-third the length; the apex is apparently strengthened 
by a straight, narrow rod that extends from the interior of the valve to the 
extreme apex, occupying about the apical .sixth of the length. 

Mesamia vitellina (Fitch). 

rPI. XXIII, fig. 7; pi. XXIX, fie. 10.) 

Length, 1.47 mm.; greatest width, 0.13 mm. Curved basal connection nar- 
row and rodlike, widens out into broader area, which is about the same width 
for entire length and toothed apically; a small preapical prominence is present 
on the ventral edge; distinctlj^ curved, tip narrowed, narrowly rounded at ex- 
treme apex, chitinization moderately heavy; strengthening rod extends caudad 
as far as fifth dorsal tooth from apex. Toothed area on dorsal edge occupies 
a little less than the apical half; teeth seventeen in number, rather small, in 



242 The Univeksity Science Bulletin. 

the general shape of a greatlj^ flattened obtuse triangle with a rounded apex, 
fairly regular in size, shape and spacing, those distad being very flat and in- 
distinct, bear small; numerous secondary teeth on both cephalic and caudal 
edges; tip notched with small teeth on both edges, not continuous around the 
tip, twelve to thirteen present between preapical prominence and apex, area 
of ducts granular in appearance, circular duct openings visible; open along 
ventral apical edge, at extreme apex, along dorsal edge of toothed area, and 
in basal portion of the valve; the two valves of the pair are joined one to the 
other by an elongate, distinct, heavily chitinized connection present on the 
dorsal edge at a point a little more than one-third the length of the valve ; the 
apex is apparently strengthened by a short, narrow rod which can be seen 
only at the extreme apex, much shorter and lighter than in M. straminca. 

Genus Scaphoideus Uhler. 

The ovipositors of two species of this genus have been examined. 
They do not resemble each other to the degree that is common be- 
tween two species of the same genus, though they have many points 
in common. In each case the ovipositor is about the same width for 
the entire length, narrows only at the apex, and bears no preapical 
prominence. The dorsal edge bears teeth along the apical two-fifths 
of its length, but these teeth differ in number, size, shape, and pos- 
session of secondary teeth in the two species, the tip bears teeth in 
one species, none in the other; the area of ducts is granular in ap- 
pearance and the duct openings are visible. The two valves of the 
pair are joined one to the other by an elongate, heavily chitinized 
connection present on the dorsal edge about midway of the valve. 

Scaphoideus scalaris Van Duzee. 

(PL XXX, fig. 1.) 

Length, 1.53 mm.; greatest width, 0.13 mm. Curved basal connection nar- 
row and rodlike, rest of valve about the same width for entire length, narrow- 
ing only at apex; beai*s no preapical prominence; slightly curved, tip narrowed, 
bluntly rounded at extreme apex, chitinization rather heavy ; strengthening rod 
extends caudad almost to second dorsal tooth from apex. Toothed area on 
dorsal edge occupies a little more than the apical two-fifths of the length; 
teeth nine in number, rather large, broadlj^ rounded, rather irregular in size, 
shape and spacing being farther apart basally; bear no secondary teeth; tip 
bears no teeth, ducts granular in appearance, circular duct openings visible; 
open along ventral apical edge, at extreme apex, along dorsal edge of toothed 
area, and irregularly in the basal region where elongate ducts are visible; the 
two valves of the pair are joined one to the other by an elongate, heavily 
chitinized connection present on the dorsal edge about midway of the valve ; 
this connection occupies a little less than a third of the entire length and bears 
the first dorsal tooth. 



Readio: Ovipositors of Cicadellid.e. 248 

Scaphoideiis immistus (Say). 

(PI. XXIII, fig. 4; pi. XXX, fig. 2.) 

Length, 1.77 mm.; greatest width, 0.11 mm. Narrow and rodliko at base, 
rest of vahe somewhat broader, about the same width for entire length, nar- 
rows caudad to ape.\; bears no preapical prominence; distinctly curved, tip 
greatly narrowed, extreme apex rounded, chitinization moderately heavy; 
strengthening rod extends caudad as far as fifth dorsal tooth from apex. 
Toothed area on dorsal edge occupies a little less than the apical two-fifths 
of the length; teeth sixteen in number, rather small, rounded, rather irregular 
in size, shape and spacing being farther apart basally; may bear secondary on 
one or both edges, small secondarj- teeth also pi-esent on margin of valve be- 
tween primary teeth, general arrangement of secondary teeth very irregular; 
tip notched with small, irregular teeth on both edges, though more distinct on 
the ventral edge, not continuous around the tip, eight to nine present on the 
ventral edge; ducts partly granular and pai'tly elongate, circular openings visi- 
ble ; open along ventral apical edge, at extreme apex, along dorsal edge of 
toothed area, and irregularly in the basal region; the two valves of the pair 
are joined one to the other by an elongate, heavily chitinized connection 
present on the dorsal edge about midway, occupies about two-fifths of length, 
bears one rounded tooth somewhat caudad of the center. 

Genus Platymetopius Burmeister. 

The ovipositors of three members of this genus have been ex- 
amined and found to be similar. In each case the basal half is 
slightly narrower than the apical half, which bears teeth along its 
dorsal edge. A preapical prominence may or may not be present. 
The primary teeth bear secondary teeth on both edges, more on the 
caudal than on the cephalic edge. In every case the tip is notched 
with small teeth on both dorsal and ventral edges; these may or 
may not be continuous around the tip. The area of ducts may or 
may not be conspicuous, when visible is granular in appearance with 
the duct openings visible. The two valves of the pair are joined 
one to the other by a rather short, chitinous connection present on 
the dorsal edge of the basal area. 

Platy77ietopius acutus (Say). 

(Pl. XXIII, fig. 5; pi. XXX, fig. 3.) 

Length, 1.36 mm.; greatest width, 0.17 mm. Curved basal connection nar- 
row and rodlike; beyond this is a somewhat wider portion extending nearly to 
midpoint, and apically the toothed area, about the same width for entire 
length, narrowing only at apex, bears an indistinct, broadly angled preapical 
prominence on the ventral edge; slightly though distinctly curved, tip nar- 
rowed and ending in a bluntly rounded extreme apex, chitinization light; 
strengthening rod extends caudad as far as fifth dorsal tooth from apex. 
Toothed area on dorsal edge occupies the apical half of the valve; teeth 
twenty-seven in number, rather large; in the greater number of teeth the basal 



244 The University Science Bulletin. 

edges of each tooth are more or less parallel and the apical edges taper, giving 
a gabled appearance to the tooth; the caudal edge is somewhat longer than 
the cephalic, fairly regular as to size, shape and spacing, though not entirely so; 
primary teeth bear small secondary teeth on caudal edge more commonly, 
and a few teeth also bear secondary teeth on the cephalic edge; tip notched 
with small teeth on both dorsal and ventral edges, continuous around the tip, 
nineteen present on ventral edge between preapical prominence and apex; 
area of ducts inconspicuous, the apices and circular openings alone being 
visible; open along ventral ai)ical edge, at extreme apex, along dorsal edge of 
toothed area, and irregularly in the basal portion of the valve; the two valves 
of the pair are joined one to the other by an indistinct, chitinous connection 
present on the dorsal edge at about a third of the length. 

Platymetopius cinereus Osborn. 

(PI. XXX, fig. 4.) 

Length, 1.13 mm.; greatest width, 0.13 mm. Curved basal connection nar- 
row and rodlike; a wider area extends almost to midpoint, a slighter wider 
apical portion occupies the rest of the valve; no preapical prominence; 
slightly curved, tip narrowed, ends in obtuse-angled point, chitinization mod- 
erately light; strengthening rod extends caudad as far as sixth dorsal tooth 
from apex. Toothed area on dorsal edge occupies a little more than the apical 
half; teeth twenty-three to twenty-four in number, moderately large, of a 
general triangular shape with the caudal side in most cases longer than the 
cephalic and with apices rounded, fairly regular in size, shape and spacing; 
bears small secondary teeth on caudal edge, three to eight in number, and a few 
of the teeth also bear a single secondary tooth on the cephalic edge; tip 
notched with small teeth on both dorsal and ventral edges, more numerous and 
distinct on ventral edge, not continuous around the tip, fourteen present on 
the ventral edge; ducts very inconspicuous, only a few being visible in the 
apex of the valve, apices of ducts and duct openings visible ; open along ventral 
apical edge, at extreme apex, along dorsal edge of toothed area, and irregu- 
larly in the basal portion of the valve; the two valves of the pair are joined 
one to the other by a poorly defined, heavily chitinized connection present on 
the dorsal edge at about one-third the length. 

Platyjnetopius frontalis Van Duzee. 

(PI. XXIII, fig. 6; pi. XXX, fig. 5.) 

Length, 1.26 mm.; greatest width, 0.16 mm. Curved basal connection nar- 
row and rodlike; a slightly wider portion extends to a point nearly midway; 
the apical portion is slightly wider, naiTOws caiidad to apex, bears a suggestion 
of a broadly rounded preapical prominence on the ventral edge; shghtly 
curved, tip narrowed, ends in a blunt, obtuse-angled point, chitinization mod- 
erately heavy, heavier than in P. acutus and P. cinereus; strengthening rod 
extends caudad as far as fifth dorsal tooth from apex. Toothed area on dorsal 
edge occupies a little more than the apical half; teeth twenty-one to twenty- 
two in number, rather large; in the greater number the basal edges of each 
tooth are parallel and the apical edges taper to a rounded apex, fairly regular 
in size, shape and spacing ; bear small secondary teeth for the most part on the 



Readio: Ovipositors of Cicadellid.e. 245 

caudal edge; tip iiotrliod with tiiuall teeth ou both dorsal and ventral edges, 
practically continuous around the tip, eighteen present on the ventral edge; 
area of ducts conspicuous, granular in appearance, circular duct openings vis- 
ible: open along ventral apical edge, along dorsal edge of toothed area, and 
irregularly in the basal area of the valve; the two valves of the pair are joined 
one to the other by a distinct, rectangular, heavily chitinized connection pres- 
ent on the doi-sal edge at about one-third the length. 

Genus Deltocephalus Burmeister. 
The u\ipositors of seven species of this genus have been examined 
and found to be similar. The ovipositor is slightly curved, gradually 
narrowed toward the apex, with apical half little if any wider than 
the basal half. The apical half bears teeth along its dorsal edge, 
usually small, rounded, and bearing a few secondary teeth. The 
apex may or may not bear small teeth on one or both edges. The 
area of ducts may or may not be conspicuous ; when visible is gran- 
ular in appearance with duct openings visible. The two valves of 
the pair are joined one to the other by a chitinous connection pres- 
ent on the dorsal edge of the basal area. 

Deltocephalus reflexus Osborn and Ball. 

(PI. XXX, fig. 6.) 

Length, 1.18 mm.; greatest width, 0.09 mm. About the same width for en- 
tire length; tapers gradually caudad to tip; no preapical prominence; slightly 
but distinctly curved, tip gradually narrowed, extreme apex sharply pointed, 
chitinization light; strengthening rod extends caudad as far as tenth doi"sal 
tooth from tip, nearly meeting dorsal edge. Toothed area on dorsal edge oc- 
cupies somewhat less than the apical half of the valve; teeth about forty in 
number, rather small, of various sizes and shapes, presenting an irregular, 
crenulate margin; a few of the larger teeth bear secondary teeth; the extreme 
tip bears no teeth ; ducts inconspicuous, invisible except for apices and circular 
openings; open along ventral apical edge, along dorsal edge of toothed area, 
and irregularh- in the basal portion of the valve; the two valves of the pair 
are joined one to the other by a poorly defined, rather elongate, heavily chit- 
inized connection present on the dorsal edge a little past a third of the length. 

Deltocephalus weedi Van Duzee. 

(PI. XXX, fig. 7.) 

Length, 1.05 mm.; greatest width, 0.13 mm. Narrow and rodlike at base; 
a somewhat wider, rather heavily chitinized area extends almost to midpoint ; 
the apical half is occupied by a still wider, less heavily chitinized area which 
tapers to apex; preapical prominence wanting; slightly curved, tip greatly 
narrowed, extreme apex rounded, chitinization moderately heavy; heavier than 
in D. reflexus; strengthening rod extends caudad as far as or beyond last dor- 
sal tooth. Toothed area on dorsal edge occupies a little more than the apical 
half of the valve; about thirteen large primary teeth, these are comparatively 



246 The University Science Bulletin. 

small, rounded, fairly regular in size, shape and spacing; bear a few small 
secondary teeth on caudal edge ; margin of valve between teeth is also notched 
with, small secondary teeth; tip notched with small teeth on both dorsal and 
ventral edges, not continuous around the tip, nine present on ventral edge; 
area of ducts conspicuous, granular in appearance, circular duct openings vis- 
ible; open along ventral apical edge, at extreme apex, along dorsal edge of 
toothed area, and irregularly in the basal area where elongate ducts are vis- 
ible; the two valves of the pair are joined one to the other by an elongate, 
distinct, heavily chitinized connection present on the dorsal edge about a third 
of the length. 

Deltocephalus inimicus (Say). 

(PI. XXIII, fig. 8; pi. XXX, fig. 8.) 

Length, 1.1 mm.; greatest width, 0.11 mm. About the same width for en- 
tire length, narrows caudad to apex, preapical prominence wanting; slightly 
curved; tip greatly narrowed, extreme apex rounded, chitinization moderately 
heavy; strengthening rod extends caudad "as far as last dorsal primary tooth. 
Toothed area on dorsal edge occupies apical half; primary teeth thirteen in 
number, small, rounded, rather regular as to size, shape and spacing; bear a 
few small secondary teeth on caudal edge; margin of valve between primary 
teeth is also notched with small teeth; ventral edge of tip is notched with 
small teeth, dorsal edge devoid of teeth at apex, fifteen present on ventral 
edge; area of ducts somewhat granular, though elongate ducts are present, 
circular duct openings visible; open along ventral apical edge, at extreme 
apex, along dorsal edge of toothed area and irregularly in the basal region ; the 
two valves of the pair are joined one to the other by an elongate, distinct, 
heavily chitinized connection present on the dorsal edge at about one-third 
the length. 

Deltocephalus jlavicosta Stal. 

(PI. XXIII, fig. 9; pi. XXX, fig. 9.) 

Length, 1 mm.; greatest width, 0.1 mm. Apical portion bearing teeth 
occupies a little more than one-half the length, is slightly wider than the 
basal portion, narrows caudad to apex; no preapical prominence; slightly 
curved, tip narrowed, extreme apex bluntly rounded, chitinization moderately 
heavy; strengthening rod extends caudad as far as last primary tooth. 
Toothed area on dorsal edge occupies a little more than the apical half; 
teeth twelve to thirteen in number, small, rounded, rather regular in size, 
shape and spacing; bear a few small secondary teeth along the caudal edge; 
margin of valve between primary teeth is also notched with small secondary 
teeth; tip notched with small teeth on ventral edge, dorsal edge devoid of 
teeth at apex, eleven present on ventral edge; area of ducts somewhat granu- 
lar in appearance, though elongate ducts are visible; open along ventral apical 
edge, at extreme apex, along dorsal edge of toothed area, and irregularly in 
the basal area; the two valves of the pair are joined one to the other by an 
elongate, curved, heavily chitinized connection present on the dorsal edge at 
about one-fourth the length. 



Readio: Ovipositors of Cicadellid.e. 247 

Deltocephalus debilis Uhler. 

(PI. XXX, fig. 10.) 

Length, 1.33 mm.; greatest width, 0.15 mm. About the same width for en- 
tire length beyond curved basal connection, narrows caudad to apex; pre- 
apical prominence wanting; distinctly curved, tip gradually narrowed, extreme 
apex rovmded, chitinization moderatelj^ heavy; strengthening rod extends 
caudad as far as fourth dorsal tooth from apex. Toothed area on dorsal edge 
occupies apical two-fifths of length; primary teeth thirteen to fourteen in 
number, small, usually rounded, irregular as to size, shape and spacing; for 
the most part bear no secondary teeth, though a few irregular secondary 
teeth are present ; the margin of the valve between the primary teeth is 
notched with small, irregular teeth; the tip is slightly and irregularly rough- 
ened, but bears no distinct teeth; area of ducts conspicuous, granular in ap- 
pearance, though elongate ducts are visible; open along ventral apical edge 
(five), at extreme apex (one), along dorsal edge of toothed area, and in the 
basal portion of the valve; the two valves of the pair are joined one to the 
other by a distinct, elongate, slightly curved, heavily chitinized connection 
present on the dorsal edge of the basal area, occupying more than one-third 
the entire length. 

Deltocephalus parvulus Gillette. 

(PI. XXX, fig. 11.) 

Length, 0.87 mm.; greatest w^idth, 0.08 mm. About the same width for 
entire length beA^ond curved basal connection, tapers gradually caudad to 
apex; preapical prominence wanting; only slightly curved, tip narrowed, ends 
in sharp point, chitinization light ; strengthening rod extends caudad almost 
to last dorsal primary tooth. Toothed area on dorsal edge occupies a little less 
than the apical half; primary teeth nine to ten in number, very small, rounded, 
fairly regular in size, shape and spacing, in most cases bear a few secondary 
teeth; margin of valve between primary teeth also bears small, regular sec- 
ondary teeth, which point forward; tip devoid of teeth; ducts inconspicuous, 
their apices and circular openings alone being visible; open along ventral 
apical edge, at extreme apex, along dorsal edge of toothed area, and in the 
basal area where elongate ducts are visible; the two valves of the pair are 
joined one to the other by an elongate, cui^ved, distinct, heavily chitinized 
connection present on the dorsal edge of the basal area and occupying about 
one-fifth of the entire length. 

Deltocephalus collinus Boheman. 

(PI. XXX, fig. 12.) 

Length, 1.3 mm.; greatest width, 0.13 mm. Narrow and rodlike at base; 
beyond base a broader and more heavily chitinized portion extends to mid- 
point; apical portion is still wider, less heavily chitinized, tapers caudad api- 
cally; preapical prominence wanting; slightlj' curved, tip greatly nan-owed, 
extreme apex nan-owly rounded, rather blunt, chitinization moderately light; 
strengthening rod extends caudad as far as seventh primary tooth from apex. 
Toothed area on dorsal edge occupies a little less than the apical half; about 
seventeen primary teeth present, these are small, somewhat rounded, very irreg- 



248 The University Science Bulletin. 

ular as to size, shape and spacing and may or may not bear a few small second- 
ary teeth on the caudal edge; margin of valve between primary teeth notched 
with small, irregular teeth, especially caudad of apex; tip devoid of teeth; area 
of ducts conspicuous, granular in appearance, though elongate ducts are visible, 
circular duct openings visible; open along ventral apical edge (five), at ex- 
treme apex (one), along dorsal edge of toothed area, and irregularly in the 
basal area; the two valves of the pair are joined one to the other by a distinct, 
elongate, slightly curved, heavily chitinized connection present on the dorsal 
edge, occupying about one-fifth of entire length. 

Aconura argentiolus (Uhler) . 

(PI. XXXI, fig. 9.) 

Length, 1.77 mm.; greatest width, 0.11 mm. About the same width for en- 
tire length, tapers caudad to apex ; bears no i^reapical prominence ; slightly 
curved, tip narrowed, rather blunt, obtuse-angled point, chitinization very 
light; strengthening rod extends caudad to a point between last and next to 
last dorsal tooth. Toothed area on the dorsal edge occupies a little more than 
the apical half; teeth twenty-two in number, very small, in the general shape 
of a greatly flattened obtuse triangle with the caudal edge longer than the 
cephalic, rather irregular in size, shape and spacing; bear many small, regular 
secondary teeth along both edges, more numerous on the caudal edge, continu- 
ous along entire dorsal margin of valve toward apex; tip notched with small 
regular teeth on both dorsal and ventral edges, not continuous around the tip, 
teeth on ventral edge slightly larger than those or dorsal ; areas of ducts incon- 
spicuous; ducts invisible except for apices and circular openings; open along 
ventral apical edge, in extreme apex, along dorsal edge of toothed area, and 
irregularly in the basal portion of the valve. 

N'ephotettix cvrtipennis (Gillette and Baker). 

(PI. XXXI, fig. 10.) 

Length, 2.5 mm.; greatest width, 0.25 mm. Apical toothed half slightly 
wider than basal portion, tapers caudad to apex; bears an indistinct, broadly 
rounded preapical prominence on ventral edge; only slightly curved, tip nar- 
rowed, ends in obtuse-angled point, chitinization moderately heavy; strength- 
ening rod extends caudad beyond last distinct primary tooth. Toothed area 
on dorsal edge occupies the apical half; teeth twenty-four to twenty-five in 
number, of medium size, in the general shape of a very flat, obtuse triangle 
with the caudal side longer than the cephalic, fairly regular in size, shape and 
spacing; bear small, regular, secondary teeth on both edges, continuous around 
the apex of each tooth, seven to fifteen on caudal edge and one to four on 
cephalic edge; tip notched with small teeth on both dorsal and ventral edges, 
not continuous around the tip, about 55 on ventral edge, area of ducts con- 
spicuous, granular in appearance though elongate ducts are visible, open along 
ventral apical edge (five), at extreme apex (one), along dorsal edge of toothed 
area, and a few in the basal region; the two valves of the pair are joined one 
to the other by an elongate, narrow, heavily chitinized connection extending 
the entire length on the basal area. 



Readio: Ovipositors of Cicadellid^. 249 

Dnotura gammaroides (Van Duzee). 

(PI. XXIII, fig. 10; pi. XXXI, fig. 11.) 

Length, 1.8 mm.; greatest width, 0.21 mm. Apical half slightly wider than 
basal half, tapere caudad to apex; bears only a suggestion of a preapical promi- 
nence on the ventral edge; slightly curved, tip narrowed with extreme apex 
narrowly rounded, chitinization moderately heavy; strengthening rod extends 
caudad beyond last distinct dorsal tooth. Toothed area on dorsal edge occu- 
pies the apical half; teeth twenty in number, of medium size, in the general 
shape of a flat, obtuse triangle with the caudal edge longer than the cephalic, 
rather regular in size, shape and spacing; bear small, regular secondary teeth on 
both edges, continuous around the apex of each tooth, seven to eighteen on 
caudal edge, two to se\en on ventral edge; tip notched with small teeth on 
both dorsal and ventral edges, practically continuous around the tip though 
greatly reduced at extreme apex; those on ventral edge are larger and more 
distinct, about fifty-four present on ventral edge; area of ducts conspicuous, 
granular in appearance, duct openings visible; open along ventral apical edge 
(five), at extreme apex (one), along dorsal edge of toothed area, and in-egu- 
larly in the basal area, the two A'alves of the pair are joined one to the other 
by an elongate, narrow, heavily chitinized connection present on the dorsal 
edge of the basal area for its entire length. 

Tliis ovipositor is very similar in general appearance to that of Nephotettix 
curtipennis. 

Genus Euscelis BruUe. 

The ovipositors of six species of this genus have been examined 
and a wide variety of forms found within the genus. Euscelis exiti- 
osus (Uhler) and Euscelis striolus (Fallen), both in subgenus Athy- . 
sanus, are not similar in any except their grosser details. Euscelis 
anthracinus (Van Duzee), in subgenus Euscelis, and Euscelis comma 
(Van Duzee), in subgenus Conomellus, are similar one to the other 
but not to any other species of the genus examined. Euscelis cur- 
tesii (Fitch) and Euscelis bicolor (Van Duzee), both in subgenus 
Stirellus, are very similar one to the other but not to any other spe- 
cies of the genus examined. 

The ovipositors of the subgenus Stirellus are somewhat narrower 
in the basal half than in the apical half, which tapers to the apex 
and bears teeth along its dorsal edge. The teeth differ in number 
and in the number of secondary teeth they bear. The apex is 
notched with small teeth present on the ventral edge only. The 
area of ducts is conspicuous, granular in appearance, with the duct 
openings visible. The two valves of the pair are joined one to the 
other by an elongate, chitinized connection present on the dorsal 
edge of the basal area. 



250 The University Science Bulletin. 

Euscelis exitiosus (Uhler). 

(PI. XXIII, fig. 11; pi. XXXI, fig. 1.) 

Length, 2 mm.; greatest width, 0.2 mm. Apical portion, which occupies 
two-thirds of length, is slightly wider than basal portion, tapers caudad to 
apex; bears a distinct, angled, preapical prominence on ventral edge; only 
slightly curved beyond curved basal attachment, rather abruptly narrowed at 
tip, ends in sharp point, chitinization moderately heavy; strengthening rod ex- 
tends caudad as far as last doreal tooth. Toothed area on dorsal edge occupies 
apical two-thirds of length; teeth thirty-four to thirty-five in number, rather 
small, of a general triangular shape, with the caudal edge longer than the 
cephalic and the apex rounded, fairly regular in size, shape and spacing; bear 
small secondary teeth on both edges, not continuous around the apex of each 
tooth, eight to eighteen on caudal edge, two to four on cephalic edge; tip 
notched with small, regular teeth on both dorsal and ventral edges, practically 
continuous around the tip, about eighty on ventral edge between preapical 
prominence and extreme apex; area of ducts conspicuous, granular in appear- 
ance; open along ventral apical edge, along the dorsal edge of the toothed 
area, one in each tooth, and a very few in the basal area; the two valves of 
the pair are joined one to the other by a poorly defined connection which is 
slightly more heavily chitinized than the rest of the basal area, upon whose 
dorsal edge it is located. 

Euscelis striolus (Fallen). 

(PI. XXIII, fig. 12; PL XXXI, fig. 2.) 

Length, LI mm.; greatest width, 0.16 mm. Apical portion occupying two- 
thirds of length but little wider than basal portion, narrows caudad to tip; 
bears no preapical prominence; only slightly cuiwed, tip abruptly narrowed, 
ends in obtuse-angled point, chitinization moderately heavy; strengthening 
rod extends caudad as far as second dorsal tooth from apex. Toothed area on 
dorsal edge occupies apical two-thirds; teeth eleven to twelve in number, 
rather large, of a general triangular shape with the apices broadly rounded, 
fairly regular in size and shape, irregular in spacing; bear secondary teeth of 
various sizes and shapes on both edges, three to nine on caudal edge and one 
to four on cephalic edge; general arrangement of secondary teeth is very ir- 
regular; tip bears a few irregular teeth on the dorsal edge only, there is a 
slight irregularity of the margin on the ventral apical edge; area of ducts con- 
spicuous, granular in appearance, circular duct openings easily visible; open 
along ventral apical edge, in extreme apex, along dorsal edge of toothed area, 
and in basal region where many elongate ducts are visible; the two valves of 
the pair are joined one to the other by an elongate, distinct, chitinized con- 
nection present on the dorsal edge of the basal area. 

Euscelis anthracinus (Van Duzee). 

(PL XXXI, fig. 3.) 

Length, L4 mm.; greatest width, 0.18 mm. About the same width for en- 
tire length beyond curved basal connection; preapical prominence wanting; 
slightly curved, tip narrowed by broadly rounded ventral edge, extreme apex 
bluntly rounded, cliitinization rather heavy; strengthening rod extends caudad 
as far as last dorsal tooth. Toothed area on dorsal edge occupies apical half; 



Readio: Ovipositors of Cicadellid^. 251 

teeth nine to ten in number, veiy small, rounded, fairly regular as to size, 
shape and spacing, being farther apart basally; bear a few small secondaiy 
teeth on caudal edge, margin of valve between teeth also notched with small, 
numerous secondary teeth; tip notched with small teeth on both dorsal and 
\entral edges, not continuous around the tip, about fifteen present on the 
ventral edge; area of ducts conspicuous, granular in appearance, though elon- 
gate ducts are visible; duct openings visible; open along ventral apical edge, 
at extreme apex, along dorsal edge of toothed area, and in basal region, where 
many elongate ducts are visible; the two valves of the pair are joined one to 
the other by a distinct, elongate, curved, heavily chitinized connection present 
on the dorsal edge of the basal area; this connection bears a rounded promi- 
nence toward the caudal end. 

Euscelis comma (Van Duzee). 

(PI. XXIII, fig. 13; pi. XXXI, fig. 4.) 

Length, 1.5 mm.; greatest width, 0.22 mm. About the same width for entire 
length beyond curved basal connection, tapers caudad to apex; no preapical 
prominence; only slightly curved, tip narrowed by broadly curving ventral 
edge, extreme apex narrowly rounded, chitinization rather heavy; strengthen- 
ing rod extends caudad to within a short distance of apex. Toothed area 
on dorsal edge occupies a little less than the apical half; teeth numerous, 
small, roimded, veiy irregular as to size, shape and spacing and present an 
irregularlj' crenulate margin; tip bears no distinct teeth but is irregularly 
roughened; area of ducts conspicuous, granular in appearance, with many 
elongate ducts visible; open along ventral apical edge, at extreme apex, along 
doi-sal edge of toothed area, and in the basal region where many elongate ducts 
are visible; the two valves of the pair are joined one to the other by an 
elongate, distinct, heavily chitinized connection present on the dorsal edge of 
the basal area; it is a little less than half the entire length and at the point 
of greatest width is about one-half the width of the valve; bears an indistinct, 
broadly rounded prominence a little caudad of the midpoint. 

Euscelis curtfsii (Fitch). 

(PI. XXXI, fig. 5.) 

Length, 1.1 mm.; greatest width, 0.12 mm. Apical half slightly wider than 
basal half, bears no preapical prominence ; only slightly curved beyond curved 
basal attachment, gradually narrowed, ends in narrowly rounded extreme apex, 
chitinization moderatelj' light; strengthening rod extends caudad as far as 
next to last dorsal tooth. Toothed area on dorsal edge occupies the apical 
half; teeth fourteen to sixteen in number, small, rounded, fairly regular in size, 
shape and spacing, though not entirely so; bear a few secondar.v teeth on the 
caudal edge and an occasional secondary tooth on the cephalic edge; tip notched 
with small teeth on ventral edge onh', dorsal edge devoid of teeth at extreme 
apex, ten present on ventral edge; area of ducts conspicuous, granular in ap- 
pearance, openings visible; open along ventral apical edge, at extreme apex, 
along dorsal edge of toothed area, and in basal region where elongate ducts are 
visible; the two valves of the pair are joined one to the other by a distinct, 
elongate, slightly curv-ed, chitinized connection present on the dorsal edge of 
the basal area. 



252 The University Science Bulletin. 

Euscelis bicolor (Van Duzee). 

(PI. XXXI. fig. 6; pi. XXIII, fig. 14.) 

Length, 0.92 mm.; greatest width, 0.11 mm. Apical portion shghtly wider 
than basal portion, tapers caiidad to apex; no preapical prominence; distinctly 
curved, tip gradually narrowed, ends in naiTowly rounded extreme apex, 
chitinization rather light; strengthenuig rod extends caudad as far as next 
to last dorsal tooth. Toothed area on dorsal edge occupies apical two-fifths 
of length; teeth twelve to thirteen in number, small, rounded, fairly regular 
in size, shape and spacing; bear a few small secondaiy teeth on caudal edge, 
margin of valve also notched with small secondar>' teeth; tip notched with 
small teeth on ventral edge only, eight to ten in number; area of ducts con- 
spicuous, granular in appearance, duct openings visible; open along ventral 
apical edge, at extreme -apex, along dorsal edge of toothed area, and in basal 
region, where elongate ducts are visible; the two valves of the pair are joined 
one to the other by an elongate connection only slightly more* heavily 
chitinized than the rest of the basal area, occupying about one-sixth of the 
entire length. 

Genus Eutettix Van Duzee. 

The ovipositors of two species of this genus have been examined 
and found to be generally similar, though possessing many differ- 
ences. In each case the basal half is somewhat narrower than the 
apical half, which narrows toward the apex and bears teeth along 
its dorsal edge. The teeth vary in number, size, shape, and posses- 
sion of secondary teeth in the two species. The tip is notched with 
small teeth on one or both edges. The area of ducts is conspicuous, 
granular in appearance, with the duct openings visible. The two 
valves of the pair are joined one to the other by a chitinous con- 
nection present on the dorsal edge of the basal area. 

Eutettix cinctus Osborn and Ball. 

(PI. XXIV, fig. 1; pi. XXXI, fig. 8.) 

Length, 1.5 mm.; greatest width, 0.17 mm. Apical portion but slightly wider 
than basal portion, narrowed caudad toward apex; no preapical prominence; 
distinctly curved, tip narrowed, extreme apex broadly rounded, chitinization 
moderately heavy; strengthening rod extends caudad as far as fourth dorsal 
tooth from apex. Toothed area on dorsal edge occupies apical half of valve; 
teeth fourteen in number, of medium size, rounded, fairly regular in size, shape 
and spacing; bear secondary teeth, one to four on caudal edge; cephalic edge 
may also bear a single secondary tooth ; margin of valve is notched with small 
secondary teeth between primary teeth; a few faintly visible teeth are present 
on the ventral edge of the tip, the dorsal edge is devoid of teeth at the extreme 
apex, about nine present on,; the ventral edge; area of ducts conspicuous, 
granular in appearance, duct openings easily visible; open along ventral apical 
edge (seventeen to nineteen), at extreme apex (three), along dorsal edge 
of toothed area, and in the basal region where elongate ducts are visible; the 



Readio: Ovipositors of Cicadellid^, 253 

two valves of tho pair arc joined one to the other by a distinct, elongate, 
curved, heavily chitinized connection present on the dorsal edge of the basal 
area; bears a distinct, rounded prominence about midway. 

Eutettix strobi (Fitch). 

(PI. XXIV, fig. 2; pi. XXXI, fig. 7.) 

Length. 1.3 ami.; greatest width. 0.17 mm. Apical portion somewhat wider 
than basal portion, narrows caudad to apex; preapical prominence wanting; 
slightly curved, tip narrowed by curving dorsal edge, ends in broadly rounded 
extreme apex, chitinization medium; strengthening rod extends caudad as 
far as fifth doi-sal tooth from apex. Toothed area on dorsal edge occupies a 
little more than the apical half; teeth twenty-one to twenty-four in number, 
large, somewhat iiregidarly rounded with the caudal edge longer and more 
gently sloping than the cephalic edge; bear small secondary teeth on both 
edges, three to seven on caudal edge and one to three on cephalic edge; tip 
is notched with small teeth on both dorsal and ventral edges, not continuous 
around the tip, fifteen to nineteen on ventral edge; area of ducts conspicuous, 
granular in appearance, duct openings easily visible; open along ventral apical 
edge (fourteen), at extreme apex (one), along dorsal edge of toothed area, and 
in basal region where elongate ducts are visible; the two valves of the pair are 
joined one to the other by a rather short, distinct, heavily chitinized con- 
nection present on the dorsal edge in the basal region. 

Genus Phlepsils Fieber. 

The ovipositors of three species of this genus have been examined 
and found to be generally similar. The ovipositor in each case is 
about the same width for entire length and tapers caudad to the 
apex. The toothed area on the dorsal edge occupies from a third 
to a half the apical length. The primary teeth vary in number, 
size, shape and number of secondary teeth in the various species. 
The tip is notched with small teeth on both edges in every case. The 
area of ducts is conspicuous, granular in appearance, with some 
eloiigate ducts visible and with duct openings visible ; the two valves 
of the pair are joined one to the other by an elongate, chitinized 
connection present on the dorsal edge of the basal area. 

Phlepsius spatulatus Van Duzee. 

(Pl. XXIV, fig. 3; pi. XXXII, fig. 10.) 

Length, 1.9 mm.; greatest width, 0.25 mm. About the same width for 
entire length, tapers caudad to apex; bears a suggestion of a preapical promi- 
nence on the ventral edge; slightly curved, narrowed by curving ventral edge, 
rather bhmtly rounded at extreme apex, chitinization heavy; strengthening rod 
extends cauded as far as fifth dorsal tooth. Toothed area on dorsal edge 
occupies apical two-fifths; teeth fifteen to eighteen in number, of medium 
size, rounded, rather irregular as to size, shape and spacing; bear a few 
irregular secondary teeth on the caudal edge; cephalic edge of some teeth 



254 The University Science Bulletin. 

also bears a single secondary tooth; margin of valve between primary teeth 
notched with small secondary teeth; tip notched with small, irregular teeth 
on both edges; area of ducts conspicuous, granular in doi-sal region, duct-like 
in ventral region, circular duct openings visible; open at ventral apical edge 
(seven), at extreme apex (one), along dorsal edge of toothed area, and in 
basal region; the two valves of the pair are joined one to the other by a 
distinct, elongate, heavily chitinized connection present on the dorsal edge of 
the basal area and occupying about one-tliird the length. 

Phlepsius excultus (Uhler). 

(PI. XXIV, fig. i; pi. XXXII, fig. 9.) 

Length, 1.8 mm.; greatest width, 0.17 mm. About the same width for entire 
length, tapers caudad to apex; bears a suggestion of a preapical prominence 
in the ventral edge, extreme apex rounded, chitinization moderately heavy; 
strengthening rod extends caudad as far as fourth dorsal tooth from apex. 
Toothed area on dorsal edge occupies a little more than the apical third of 
the length; teeth fifteen to seventeen in number, rather small, roimded, 
fairly regular in size and shape but uneven in spacing; bear a few secondary 
teeth on caudal edge, and a few primary teeth bear a single secondary tooth 
on the cephalic edge, margin of valve between primary teeth bears small 
secondary teeth; tip notched with small teeth on both dorsal and ventral edges, 
practically continuous around the tip; eight large teeth on ventral edge; these 
may be simple or may bear secondary teeth, irregular as- to arrangement; 
area of ducts conspicuous, ducts elongate for the most part though a granular 
area is present dorsally, duct openings visible; open along ventral apical edge 
(six), at extreme apex (one), along dorsal edge of toothed area, and in the 
basal region; the two valves of the pair are joined one to the other by a dis- 
tinct, elongate heavily chitinized connection present on the dorsal edge of the 
basal area, occupying more than a third of the total length, this connection 
irregularly roughened along its dorsal edge. 

Phlepsius irroratus (Say). 

fPl. XXIV, fig. 5; pi. XXXII, fig. 8.) 

Length, 1.5 mm.; greatest wddth, 0.17 mm. About the same width for 
entire length beyond curved basal attachment, narrows caudad to apex; pre- 
apical prominence wanting; slightly curved, tip gradually narrowed, narrowly 
rounded at extreme apex, chitinization moderately light; strengthening rod 
extends caudad as far as second dorsal tooth from apex. Toothed area on 
dorsal edge occupies a little less than the apical half; teeth nineteen to twenty 
in number, of medium size, in the general shape of an obtuse triangle with 
caudal edge longer than the cephalic and the apex rounded broadly, fairly 
regular as to size shape and spacing, but not entirely so ; bear a few secondary 
teeth on caudal edge and also a few bear a single secondaiy tooth on the 
cephahc edge; tip is notched with small, indistinct teeth on both dorsal and 
ventral edge, not continuous around the tip, about fifteen on ventral edge; 
area of ducts conspicuous, granular in appearance, duct openings easily 
visible; open along ventral apical edge (ten), at extreme apex (one), along 
dorsal edge of toothed area, and in the basal region where elongate ducts are 



Readio: Ovipositors of Cicadellid^. 255 

visible; the two ^■alves of the pair are joined one to the other by a heavily 
chitinized connection present on the dorsal edge of the basal area, occupying 
about a fourth of the entire length and a half of the width. 

Genus Acinopterus Van Duzec. 

The ovipositors of three species of this genus have been examined 
and found to be very similar. In each case the ovipositor is greatly 
curved, the basal portion only slightly narrower than the apical 
portion, which bears teeth along its dorsal edge. The primary teeth 
are few in number, and differ in number and in the number of sec- 
ondary teeth they bear in the various species. The margin of the 
valve between the primary teeth is notched with small teeth. The 
tip in each case is notched with small teeth on both dorsal and 
ventral edges. The area of ducts may or may not be conspicuous, 
when visible granular in appearance with duct openings visible. 
The two valves of the pair are joined one to the other by an elon- 
gate, curved, chitinous connection present on the dorsal edge of 
the basal area. 

Acinopterios acuminatus Van Duzee. 

(PI. XXIV, fig. 6; pi. XXXII, fig. 3.) 

Length, 1.5 mm.; greatest width, 0.12 mm. Apical portion only slightly 
wider than basal portion, tapers slightly caudad; bears a small preapical 
prominence on the ventral edge; greatly curved, chitinization moderately 
light; strengthening rod extends caudad as far as last or next to last dorsal 
tooth of large size. Toothed area on dorsal edge occupies somewhat less than 
the apical half; primary teeth seven in number, small, rounded, regular as to 
size and shape, unevenly .spaced; bears three to four small secondary teeth 
on the caudal edge, margin of valve between primary teeth bears numerous 
small, regular, secondary teeth; tip notched with small teeth on both dorsal 
and ventral edges, not continuous around the tip, about fifteen present on the 
ventral edge; area of ducts conspicuous, granular in appearance, circular 
duct openings visible; open along ventral apical edge (seven), at extreme 
apex (two), along dorsal edge of toothed area, and in the basal region; the 
two valves of the pair are joined one to the other by a distinct, curved, heavily 
chitinized connection present on the dorsal edge of the basal area. 

Acinopterits viridis Ball. 

(P!. XXXII, fig. 1.) 

Length, 1.7 mm.; greatest width, 0.16 mm. Apical portion only slightly 
wider than basal portion, nan-owed caudad at apex, bears no preapical promi- 
nence; greatly curved, tip narrowed, ends in rounded extreme apex, chitiniza- 
tion light; strengthening rod extends caudad almost to apex. Toothed area 
on dorsal edge occupies the apical third; only two primary teeth present, 
these located near the base of the toothed area, small, rounded, bear two to 
three small secondary teeth on their caudal edges; the margin of the valve is 
notched with many small, regular secondary teeth; tip is notched on both 



256 The University Science Bulletin, 

edges with small teeth, continuous around the tip, about ten on ventral edge; 
area of ducts inconspicuous, ducts invisible except for circular openings; open 
along ventral apical edge (six), at extreme apex (one), along dorsal edge of 
toothed area, and in basal area; the two valves of the pair are joined one to 
the other by a poorly defined, elongate connection present on the dorsal edge 
of the basal area, only slightly more heavily chitinized than the rest of the 
basal area. 

Acinopterns angulatus Lawson. 

(PI. XXXII, fig. 2.) 

Length, 1.2 mm.; greatest width, 0.12 mm. Apical portion only slightly 
wider than basal portion, narrows slightly caudad at apex; no distinct 
preapical prominence present; greatly curved, tip narrowed, extreme apex 
rounded, chitinization moderately heavy, heavier than in A. acuminatus and 
A. viridis ; strengthening rod extends caudad as far as next to last primary 
tooth. Toothed area on dorsal edge occupies a little more than the apical 
third; teeth eight in number, small, rounded, regular in size and shape, some- 
what unevenly spaced; bear secondary teeth on caudal edge, three to foiu- in 
mmiber; the margin of valve between primary teeth is notched with small, 
numerous, regular, secondaiy teeth; tip notched with small teeth on both 
dorsal and ventral edges, practically continuous around the tip," about fifteen 
on ventral edge; area of ducts conspicuous, granular in appearance, circular 
duct openings visible; open along ventral apical edge (five to six), at ex- 
treme apex (one), along dorsal edge of toothed area, and in basal region; the 
two valves of the pair are joined one to the other by a distinct, elongate, 
heavily chitinized connection present on the dorsal edge of the basal area. 

Genus Thamnotettix Zetterstedt. 

The ovipositors of two species of this genus have been examined 
and found to be generally similar. In each case the basal half is 
distinctly narrower than the apical half, which narrows caudad at 
the apex and bears teeth along its dorsal edge. The primary teeth 
differ in number, shape, and number of secondary teeth they bear 
in the two species. The tip is notched with small teeth on both 
edges. The area of ducts is conspicuous, granular in appearance, 
with the duct openings visible. The two valves of the pair are 
joined one to the other by a heavily chitinized connection present 
on the dorsal edge of the basal area. 

Thamnotettix clitellarms (Say). 

(PI.' XXXII, fig. 5.) 

Length, 1.4 mm.; greatest width, 0.18 mm. Apical portion plainly wider 
than basal portion, narrows caudad toward apex; bears a very small preapical 
prominence on ventral edge; slightly curved, tip narrowed, extreme apex 
broadly rounded, chitinization medium; strengthening rod extends caudad as 
far as fourth dorsal tooth from apex. Toothed area on dorsal edge occupies 
a little more than the apical half; teeth twenty to twenty-three in number, of 



Re.\dio: Ovipositors of Cicadellid.e. 257 

medium size, of a general triangular shape, with the caudal edge longer than 
the cephalic and the apex rounded broadly, fairly regular as to size, shape, 
and spacing, though not entirely so; bear small secondary teeth on caudal 
edge and a few also bear a single secondary tooth on the cephalic edge; tip 
notched with small teeth on both edges, not continuous around the tip, those 
on ventral edge larger and more distinct, about fourteen present on ventral 
edge ; area of ducts conspicuous, granular in appearance, circular duct openings 
visible; present along ventral apical edge (twelve), at extreme apex (one), along 
dorsal edge of toothed area, and in basal region where elongate ducts are visible; 
the two valves of the pair are joined one to the other by a distinct, heavily 
chitinized, rather short, rectangular connection present on the dorsal edge of 
the basal area. 

Thamnotettix longidus Gillette and Baker. 

(PI. XXXII, fig. 4.) 

Length, 1.3 mm.; greatest width, 0.17 mm. Apical portion distinctlj^ wider 
than basal portion, tapers caudad at apex; no preapical prominence; slightly 
curved, tip narrowed, extreme apex narrowly rounded, chitinization medium; 
strengthening rod extends caudad as far as sixth dorsal tooth from apex. 
Toothed area on dorsal edge occupies somewhat more than apical half; thirty- 
one primarj' teeth present, in the general shape of an obtuse triangle, of 
medium size, fairlj' regular as to size and shape but not entirely so, uneven in 
spacing; bear small secondary teeth on both edges, four to twelve on caudal 
edge and one to four on cephalic edge, secondary teeth continuous around the 
apex of each tooth; tip notched with small teeth on both dorsal and ventral 
edge, not continuous around the apex, eight on ventral edge; area of ducts 
conspicuous, granular in appearance, duct openings visible; open along ven- 
tral apical edge (eight), at extreme apex (one), along dorsal edge of toothed 
area (about one to each tooth), and in the basal area; the two valves of the 
pair are joined one to the other by a distinct, elongate, heavily chitinized 
connection present on the dorsal edge of the basal area, occupies less than a 
fifth of the entire length. 

Genus Chlorotettix Van Duzee. 

The ovipositors of two species of this genus have been examined 
and found to be generally similar. The basal third may or may not 
be narrower than the apical two-thirds, which tapers to the apex 
and bears teeth along its dorsal edge. The primary teeth differ in 
number, shape, and number of secondary teeth in the two species. 
The tip is notched on both edges with small teeth, not continuous 
around the tip. The area of ducts is granular in appearance and 
the duct openings are visible. The two valves of the pair are joined 
one to the other by a chitinous connection present on the dorsal edge 
of the basal area. 



258 The University Science Bulletin. 

Chlorotettix spatidatits Osborn and Ball. 

(PI. XXIV, fig. 7; pi. XXXII, fig. 6.) 

Length, 1.5 mm.; greatest width, 0.24 mm. Apical portion much wider than 
basal portion, tapers caudad at apex; preapical prominence wanting; sHghtly 
curved, tip rather abruptly narrowed, extreme apex narrowly rounded, chitin- 
ization medium; strengthening rod extends caudad as far as last primary 
tooth. Toothed area on dorsal edge occupies the apical two-thirds; teeth 
twenty-seven to twenty-eight in number, of medium size, rather iiTegular as 
to shape, some rounded, some triangular and some flat across the top; bear 
two to seven secondary teeth on the caudal edge and may or may not bear 
one to three secondary teeth on the cephalic edge; tip notched with small 
teeth on both edges, not continuous around the tip, nine to ten on ventral 
edge ; area of ducts conspicuous, granular in appearance though many elongate 
ducts are visible, duct openings visible; open along ventral apical edge (eight), 
at extreme apex (one), along dorsal edge of toothed area, and in basal area; 
the two valves of the pair are joined one to the other by an elongate, rectangu- 
lar, chitinized connection present on the dorsal edge of the basal area, only 
slightly more heavily chitinized than the rest of the valve, occupying about 
one-sixth of entire length. 

Chlorotettix galbanatus Van Duzee. 

(PI. XXIV, fig. 8; PI. XXXII, fig. 7. J 

Length, 1.4 mm.; greatest width, 0.21 mm. About the same width for en- 
tire length, apical portion very slightly narrower than basal portion, tapers 
caudad to apex; no preapical prominence; slightly cun^ed, tip gradually nar- 
rowed, extreme apex bluntly rounded, chitinization medium; strengthening rod 
extends caudad as far as third dorsal tooth from tip. Toothed area on dorsal 
edge occupies somewhat less than the apical two-thirds; teeth twenty-eight in 
number, of medium size, some rounded, others of a general triangular shape, 
evenly spaced; bear secondary teeth, two to eight on caudal edge and an oc- 
casional single secondary tooth on the cephalic edge; tip notched with small 
teeth on both edges, not continuous around the tip, ten present on ventral 
edge; area of ducts conspicuous, granular in appearance, circular duct open- 
ings visible; open along ventral apical edge (thirteen), at extreme apex (one), 
along dorsal edge of toothed area, and in the basal region, where elongate 
ducts are visible; the two valves of the pair are joined one to the other by a 
distinct, curved, heavily chitinized connection present on the dorsal edge of 
the basal area, occupying a little less than a fourth of the entire length. 

Jassus olitorius Say. 

(PI. XXXIII, fig. 2.) 

Length, 3.7 mm.; greatest width, 0.2 mm. Very long, narrow and rodlike, 
apical portion which bears teeth only slightly if any wider than basal rod,, 
tapers only at apex; no distinct preapical prominence present; distinctly 
curved, tip narrowed, rounded at extreme apex, chitinization moderately 
heavy ; strengthening rod extends caudad as far as sixth dorsal tooth. Toothed 
area on dorsal edge occupies about the apical fourth; eleven teeth present on 
each valve with a large median tooth present between teeth one and two; 



Readio: Ovipositors of Cicadellid^. 259 

apical teeth veiy small and broadly rounded, distal teeth larger and more 
sharply pointed; bear no secondary teeth; tip bears no teeth; area of ducts 
conspicuous, ducts elongate, rather straight, circular openings visible; open 
along ventral apical edge (twenty-two), at extreme apex (one), along dorsal 
edge of toothed area, very noticeably in teeth three and four, in each of which 
three to four ducts open, and in the basal rod; the two valves of the pair are 
joined one to the other by a distinct, elongate, narrow, heavily chitinized con- 
nection present on the dorsal edge of the basal rod. 

Tinobregnius vittatus Van Duzee. 

(PI. XXXIII, fig. 1.) 

Length, 2.7 mm.; greatest width, 0.19 mm. Long, narrow and rodlike, apical 
toothed portion only shghtly wider than basal rod. tapers caudad at apex; no 
preapical prominence; distinctly curved, tip evenly naiTOwed, extreme apex 
rounded, chitinization moderately heavy, heavier than in Jasms olitorius; 
strengthening rod extends caudad as far as eighth dorsal tooth. Toothed area 
on dorsal edge occupies apical third; teeth ten to eleven in number, apical 
teeth small and broadly rounded, distal teeth larger and more sharply pointed; 
bear no secondary teeth; tip bears no teeth; area of ducts conspicuous, ducts 
elongate, rather straight, duct openings visible; open along ventral apical 
edges (twelve), at extreme apex (one), along dorsal edge of toothed area, and 
in the entire length of basal rod; the two valves of the pair are joined one to 
the other by a distinct, elongate, narrow, heavily chitinized connection present 
on the dorsal edge of the basal rod, occupying about one-fifth of the length of 
the basal shaft. 

This ovipositor is very similar to that of Jassus olitorius. 

Cicadula punctifrons var. repleta Fieber. 

(PI. XXIV, fig. 9; pi. XXXIII, fig. 3.) 

Length, 2.2 mm.; greatest width, 0.17 mm. About the same width for entire 
length, narrows ■ caudad at apex; no preapical prominence; slightly curved, tip 
narrowed by curving ventral edge, bears a finely toothed prominence on the 
dorsal edge, extreme apex narrowly rounded, chitinization very light; strength- 
ening rod extends caudad as far as fourth dorsal tooth from apex. Toothed 
area on doi"sal edge occupies the apical two-fifths; teeth fifteen in number, 
rather large, rounded, regular as to size, shape and spacing; bear a few fine 
secondary teeth on both edges, continuous around the apices of the primary 
teeth, those on caudal edge larger and more distinct than those on ventral 
edge, two to eight present on the ventral edge; there are fine, radiating hues 
extending from the interior of each tooth to the margin; the tip is notched 
with small teeth on both edges, not continuous around the tip, twenty-five on 
ventral edge; area of ducts inconspicuous, ducts invisible except for circular 
openings, which are easily visible; open along ventral apical edge (five), at 
extreme apex (one), along dorsal edge of toothed area, and in the basal area; 
the two valves of the pair are joined one to the other by a poorly defined, 
elongate, hghtly chitinized connection present on the dorsal edge of the basal 
area, occupies about one-fourth the entire length. 



260 The University Science Bulletin, 

Genus Balclutha Kirkaldy. 

The ovipositors of two species of this genus have been examined 
and found to be similar. The basal portion is somewhat narrower 
than the apical portion. The point of greatest width is between 
three-fourths and four-fifths the length of the valve, beyond which 
the valve tapers to a very narrow apex, sharply pointed or narrowly 
rounded. The valve bears only very small teeth located at the apex. 
The area of ducts may or may not be conspicuous, is granular in ap- 
pearance with the duct openings visible. The two valves of the pair 
are joined one to the other by an elongate, lightly chitinized connec- 
tion present on the dorsal edge of the basal area. 

Balclutha punctata (Thunberg). 

(PI. XXIV, fig. 10; pi. XXXIII, fig. 4.) 

Length, 0.88 mm.; greatest width. 0.1 mm. Apical portion somewhat wider 
than basal portion, point of greatest width about three-fomtlis the length, tapers 
caudad from this point to the apex; distinctly curved, tip greatly narrowed, 
extreme apex narrowly rounded, chitinization very light; strengthening rod 
extends caudad to within a short distance of the apex. The valve is toothed 
only on its dorsal apical edge for about one-sixth the length, teeth numerous, 
very small, regular; area of ducts inconspicuous, faintly granular in appear- 
ance, duct openings visible; open along ventral apical edge, at extreme apex, 
along dorsal apical edge, and in the basal region; the two valves of the pair are 
joined one to the other by a lightly chitinized, elongate, narrow connection 
present on the dorsal edge of the basal area. 

Balclutha impicta (Van Duzee). 

(PL XXIV, fig. U; pi. XXXIII, fig. 5.) 

Length, 0.88 mm.; greatest width, 0.09 mm. Apical portio'n slightly wider 
than basal portion, point of greatest width about four-fifths the length; only 
slightly curved, tip greatly narrowed, extreme apex very sharply pointed, chi- 
tinization light, though heavier than in B. punctata; strengthening rod extends 
caudad almost to apex. The dorsal edge is toothed only for the apical sixth of 
its length; teeth numerous, very small and regular; ventral edge also bears a 
few small teeth at the apex, farther apart than those on the dorsal edge, about 
ten in number; area of ducts conspicuous, granular in appearance, openings 
visible; open along ventral apical edge, at extreme apex, along dorsal apical 
edge, and in the basal area; the two valves of the pair are joined one to the 
other by a distinct, elongate, narrow connection present on the dorsal edge of 
the basal area. 

Eugnathodus abdominalis (Van Duzee). 

(PI. XXIV, fig. 12; pi. XXXIII, fig. 6.) 

Length, 0.72 mm.; greatest width, 0.09 mm. Apical half slightly wider than 
basal half, point of greatest width is about three-fourths the length, beyond 
this point the valve tapers to the apex; distinctly curved, tip greatly but 
unevenly narrowed, extreme apex very sharply pointed, chitinization very 



Rkadu): Ovipositors ok C'kadkllid.io. 'ilil 

lifjhf ; .^trrnst honing rod extends caiidad aliiiosl to ajirx. The toothed area on 
the dorsal edge oecuijie-s only the narrowed portion of the ai)ex; these teeth 
are very small, regular, nuintM'ous; the ventral edge bears no teeth at the apex, 
but somewhat hack from the apex on the widened portion the \entral edge is 
notehcd with many fine, indistinct teeth; area of ducts conspicuous, granular 
in ajiiiearance. duct openings \ isihie ; open along ventral apical edge, at ex- 
treme ajicx, along dorsal ajiical edge, and in basal region; there is no evidence 
of a chitinous connection between the two valves of the i)air. 

This ovipositor is similar in appearance to the ovipositors of the genus 
Bnlrlnlha examined. 

Tribe Typhlocybini ( Kirschbaum I . 
Dikraneura abnorniis (Walsh). 

(PI. XXIV, fig. IS: pi. XXXIII, i\g. 7.) 

Length. 0.7 nun.; greatest width, 0.08 mm. Basal half narrow and rodlike, 
apical half wider, flat, toothed, tapers caudad to apex; basal portion greatly 
cm'ved. ajucal portion only slightly ciu'ved; tip greatly narrowed, extreme apex 
narrowly rounded, chitinization moderately light; strengthening rod extends 
caudad as far as sixth dorsal tooth from apex. Toothed area on dorsal edge 
occupies the apical half; the two valves of the pair are not identical as to 
teeth, the one having few and the other many; the one having more bears 
twenty-five primar.v teeth, the.se rather small, in the general shape of an obtuse 
triangle, fairly regular as to size and shape, much reduced in size apically; 
bear secondary teeth along the caudal edges, three to six in number; the tip is 
notched with small teeth on both edges, not continuous around the tip. four on 
the ventral edge; area of ducts inconspicuous, ducts visible, elongate, rather 
few in number; \isible openings present only along dorsal edge of toothed area 
and in basiil area; no distinct chitinous connection between the two valves 
])resent. 

(tonus Empoasc.\ Walsh. 

The ovipositors of fiv? species of this genus have been examined 
and found to be very similar. In this genus the two valves of the 
pair are not identical, but differ in length, width, and size and num- 
ber of teeth. The shorter, narrower valve bears many very small, 
regular teeth along its dorsal edge. The longer, broader valve bears 
comparatively few large teeth, which in turn bear small- secondary 
teeth. Neither valve in any of the species examined bears teeth for 
more than the apical fifth of its length. Except in one species the 
tip of the valve is notched with small teeth. The area of ducts may 
or may not be conspicuous; ducts elongate, few in number. 

Empoasca trifasciata Ctillette. 

(PI. XXXIII, fig. HI.) 

Length. 0.8 mm.; greatest width. 0.08 nun. Narrow and rodlike, apical 
toothed i)ortion onlj- slightly wider than basal shaft; di.stinctly curved, tip nar- 
rowed, extreme apex narrowly rounded, chitinization moderately light; 
strengthening rod extends caudad as far as second dorsal tooth. Toothed area 



262 The University Science Bulletin. 

on dorsal edge occupies the apical fifth of the valve; the two valves of the pair 
differ as to number and arrangement of teeth ; the shorter, narrower valve bears 
many small, regular teeth along its dorsal edge; the longer, broader valve bears 
thirteen large primaiy teeth along its dorsal edge, these rounded, regular, and 
bear one to three secondary teeth along their caudal edges; the tip is notched 
with small teeth on both dorsal and ventral edges, not continuous around the 
tip, seven on \entral edge of valve bearing small teeth, eleven on ventral edge 
of valve bearing large teeth; area of ducts conspicuous, ducts easily visible, 
elongate, rather few in number; open at apex, along dorsal edge of toothed 
area, and in basal area; no distinct chitinous connection present. 

Empoasca smaragdula (Fallen). 

(PI. XXXIII, fig. 11.) 

Length, 2 mm.; greatest width, 0.11 mm. Narrow and rodlike, about the 
same width for entire length, tapers caudad at apex; distinctly curved, tip 
narrowed, extreme apex narrowly rounded, chitinization medium; strengthen- 
ing rod extends caudad as far as third dorsal tooth. Toothed area on dorsal 
«dge occupies about the apical seventh of the length; the two valves of the 
pair are not identical; the one is shorter, narrower, and bears only very small 
teeth along its dorsal edge ; the longer, wider valve beai-s thirteen large primary 
teeth along its dorsal edge, flatly rounded, fairly regular as to size, shape and 
spacing; bear a few indistinct secondaiy teeth; the tip of the larger valve is 
notched with small, indistinct, irregular teeth on both dorsal and ventral edges, 
continuous around the apex; area of ducts conspicuous, ducts easily visible, 
elongate, few in number; open at apex and along basal shaft; the dorsal edge 
of the basal shaft is irregularly roughened; no distinct chitinovis connection 
present. 

Empoasca obtusa Walsh. 

(PI. XXIV, fig. 14: pi. XXXIII, fig. 12 ) 

Length, 0.9 mm.; greatest width, 0.05 mm. Consi.-ts of a narrow, rodlike 
basal shaft and a slightly wider, toothed apical portion which tapers caudad 
to apex; distinctly curved, tip only slightly narrowed, extreme apex broadly 
rounded, chitinization moderately light; strengthening rod extends caudad 
as far as second dorsal tooth. The two valves of the pair are not identical; 
the shorter, narrower valve bears only very small, regular teeth along its 
dorsal edge; the longer, wider valve is toothed along its apical seventh with 
seven primary teeth, these of medium size, rather flatly rounded, fairly regular 
as to size, shape and spacing, though smaller apically, and bear a few irregular, 
indistinct, secondary teeth; no distinct teeth present on tip; area of ducts 
rather conspicuous, ducts visible, few in number, elongate; open apically, 
along dorsaledge of toothed area, and along basal shaft. 

Empoasca livingstoni (iillette. 

(PI. XXXIII, fig. 14.) 

Length, 2 mm.; greatest width, 0.11 mm. Narrow and rodlike, about the 
same width for entire length, tapers caudad at apex; distinctly curved, tip 
greatly narrowed, extreme apex ends in acute-angled point, chitinization very 
light; strengthening rod extends caudad as far as fifth dorsal tooth. The two 
valves of the pair are not identical; the shorter, naiTower valve bears only 



Readio: Ovipositors of Cicadellid^. 263 

very small, lobular tooth for a short distance on its dorsal edge; the longer, 
wider \alvo bears thirteen primary teeth along the apical sixth of its dorsal 
edge, medium in size, somewhat triangular in shape, with the caudal edge 
longer than tlu^ cephalic, fairly regular in size, shape and spacing; bear small 
secondary teeth along caudal edge, three to seven in number; tip of longer 
valve notched with small teeth, continuous around the tip, thirteen on ventral 
edge; area of ducts inconsiiicuous, ducts invisible except for circular openings; 
open at ajiex. along dorsal edge of toothed area, and in basal shaft. 

Enipoasca mali (LeBaron). 

(PI. XXXIII, fig. 13.) 

Length. 0.8 mm.; greatest width, 0.03 mm. Very narrow and rodlike, apical 
toothed portion only very little wider than basal shaft; greatly curved, 
tip slightly narrowed, extreme apex narrowly rounded, chitinization light; 
strengthening rod extends caudad as far as third dorsal tooth. The two 
valves of the pair are not identical; the shorter, narrower valve bears only 
very small, regular teeth along its dorsal edge for a short distance ; the longer, 
wider valve bears eighteen primary teeth along the apical sixth of its dorsal 
edge, these of medium size, regular as to size, shape and spacing, smaller 
apically, and bear a few very fine and indistinct secondary teeth on the 
caudal edge; the tip of the longer valve is notched with small teeth on both 
edges, not continuous around the tip, about seven on the ventral edge; area 
of ducts inconspicuous, ducts invisible except for circular openings; open 
at apex and a few in the basal shaft. 

Genus Erythroneura Fitch. 

The ovipositors of two species of this genus have been examined 
and found to be similar one to the other and also to the ovipositors 
of the genus Enipoasca. In this genus the two valves of the pair 
are not identical, but differ as to length, width, and size and number 
of teeth. In one species the small, more numerous teeth are borne 
by the longer, wider valve; in the other species the condition is 
reversed. The teeth do not occur except on the apical fourth of the 
valve. The area of ducts is inconspicuous, ducts when visible elon- 
gate, duct openings visible. 

Erythroneura tricincta Fitch. 

(PI. XXXIII, fig. 8.) 

Length, L2 mm.; greatest width, 0.09 mm. Narrow and rodlike, the apical 
toothed portion only slightly wider than the basal shaft, tapers caudad at 
apex; distinctly curved, tip narrowed, extreme apex rounded, chitinization 
very light ; strengthening rod extends caudad to a point about two-fifths the 
length of the toothed portion of the longer valve. The two valves of the 
pair are not identical ; the longer, wider valve bears a great many very small, 
regular teeth along its dorsal edge for the apical fourth; the shorter, narrower 
valve bears slightly larger, less numerous, regular teeth along its dorsal apical 
edge; tip notched with teeth on both edges, not continuous aroimd the tip. 



264 The University Science Bulletin. 

about twelve on the ventral edge of the longer valve; area of ducts incon- 
spicuous, ducts invisible except for openings; open in apex and along basal 
shaft. 

- Erythroneiirn vulnerata Fitch. 

(PI. XXXIII, fig. !».) 

Length, 0.8 mm.; grratett width, 0.06 mm. Narrow and rodlike, apical 
toothed portion only slightly wider than basal shaft, tapers caudad to apex; 
distinctly curved, tij) narrowed, extreme ajiex rounded, chitinization moder- 
ately light; strengthening rod extends caudad as far as third dorsal tooth. 
The two valves of the pair are not identical ; the shorter, narrower ^-alve bears 
very small teeth for a short distance along its dorsal apical edge; the longer, 
wider valve bears ten jirimary teeth along its dorsal edge, occupying the apical 
sixth, these of medium size, in the general shape of an obtuse triangle, fairly 
regular in size, shape and spacing, bear two to five small secondary teeth on 
caudal edge; tip notched with small teeth on both edges, continuous around 
apex, about ten present on ventral edge of longer valve; area of ducts in- 
conspicuous, ducts elongate; open apically and along the basal shaft. 

CONCLUSIONS. 

An examination of the descriptions and phites leads to several 
conclusions. The various subfamilies arc not distinctly set apart 
by the characters of the ovipositor. While it is true that in general 
the ovipositors of the Bythoscopina^ have regular, rounded teeth; 
those of the Cicadellinse are toothed for nearly their entire length 
and have elongate, curved ducts; those of the Gyponinse are stout 
and heavily chitinized; and those of the Jassinse have a granular 
duct area and a chitinous connection between the two valves of the 
pair; yet these characters are not found in nil the members of the 
subfamily and are not exclusively found fii the subfamily. How- 
ever, closely related genera possess ovipositors which are very simi- 
lar. Examples of this similarity ai'e shown in Agalliopsis and Ace- 
ratagallia; Macropsis and Oncopsis; Oncometopia, Homalodisca and 
Aulacizes; Cicadella and Graphocephala; Nephotettix and Drio- 
tui'a; Dorycephalits and Hecalus; Helochara and Drcrculacephola; 
Jassus and Tinobregrnus; Balclutha and Eugnathodus; and Dikra- 
neura, Empoasca and Typhlocyba, all in the tribe Typhlocybini, 
which is very clearly set apart by the cluiracters of the ovipositor. 

Between the species of well-defined genera there is an indisputa- 
ble generic similarity. The cliaracters of size, chitinization and 
number of teeth vary, but the characters of general shape, and shape 
ahd arrangement of teeth seem to be constant within the genus. 
Examples of generic similarity are shown in Idiocerus, Macropsis, 
Kolla, Dneculacephala, Platymetopius, Deltocephahis, Balclutha, 



Readio: ( )\"ir()siTORS ok C'icAniiLLiii.K. 265 

Enipoasca, and inan>' ollicrs. In tlii' ;t<linitt('(lly loose and complex 
genus Eusceli.s the ovipositors show the wide range of forms that 
would be cxpeeted, and the same condition might be found in other 
genera of equal complexity. 

Penally, we find constant characters of specific value in the ovi- 
positor. The ovipositors of seventeen individuals of Cicadella hi- 
eroglyphica (Say), representing as wide a geographical range and 
as many color A'arieties as are to be found in our duplicate collec- 
tion, were examined and found to be constant in the characters 
given, which are sufficient to separate it from the other species of the 
genus examined. Several specimens each of Graphocephala coc- 
cinea (Forster) and Oncometopia lateralis (Fabricius) were exam- 
ined in the same way and their characters also found to be constant. 
Hence it can safely be concluded that characters of specific value, 
constant within the range of the species, are found in the ovipositors 
of the Cicadellidae. It is also true that these characters are accessi- 
ble to the general worker and should not be neglected by him in a 
taxonomic study in this family. 

BIBLIOGRAPHY. 

1878 — Packard: Guide to the Study of Insects. 

189S-.Marlatt : Periodical Cicada. U. S. Dept. of Agr., Bull. No. 14. n. s. 

1909 — Packard; Textbook of Entomology. 

1910— Stough: Hackberry Psylla. Kansas U. Sci. Bull., vol. V, No. 9. p. 121. 

1917 — Van Duzee: Catalogue of Heniip. of N. A. 

1918 — Xewell: Comp. Morph. of the Genitalia of In.sects. Ann. Ent. Soc. Am., 

vol. XI, No. 2. 11. 109. 
1919— Kornhauser: Sexual Characters of Thelia. Journ. of Morph., vol. 32. 

Xo. 3. p. 531. 
1919 — Walker: Structure of Orthoj). Insects. Ann. Ent. Soc. Am., vol. XII. 

Xo. 4, p. 267. 
1920— Lawson: Cicadellidae of Kansas. Kansas U. Sci. Bull., vol. XII. Xo. 1. 
1921 — Hilsman: Ovipositor of the Cicada. Thesis MS., University of Kansas. 



266 The University Science Bulletin. 



PLATE XXL 

1. Ventrul view of abdcmen of female. 

2. Dorsal view of abdomen of female. 

3. Cross section through abdomen. 

4. Bor-^al view of ^^egment nine, showing attachment of ovipositor. 

5. Ventral view of .•segment nine, showing attachment of ovipositor. 

6. Cross section through ovipositor, showing relative position of valves. 

7. Valve I, showing attachment to eighth sternum. 

8. Detached iiortion of valve I, showing attachment to ninth pleuron. 

9. Valve II (ujijier) and valve I (lower), showing attachment to ninth 
sternum. 



Readio: Ovipositors of Cicadellid^. 



267 



PLATE XXI. 




268 The University Science Bulletin. 



PLATE XXII. 

1. Idiucrtns pallidus Fitch. 

2. Agalliupsis novella (Say). 

3. Oncoinetopia undaia (Fabricius). 

4. Oncometopia lateralis (Fabricius). 

5. Homalodisca triquetra (Fabricius). 

6. Aulacizea irrorata (Fabricius). 

7. Cicadella hieroglyphica (Say). 

8. Cicadella circellata (Baker). 

9. Graphocephala ccccinea (Forster). 

10. Helochara communis Fitch. 

11. Drceculacephala mollipes (Say). 

12. Drceculacephala reticulata (S'gnoret). 

13. Kolla bifida (Say). 

14. Kolla hart a (Ball). 



PLATE XXII. 






10 




(269) 



PLATE XXIII. 

1. Gypona octo-Uncata (Say). 

2. Xerophloea viridis (Fabricius) . 

■3. Xestocephalus pulicariitx Van Duzce. 

4. Scaphoideus iministus (Say). 

5. Platymetopius acutus (Say). 

6. Platymetopius frontalis Van Duzee. 

7. Mesamia vitellina (Fitch). 

8. Deltoccphalus inimicus (Say). 

9. Deltoccphalus flavicosta Stal. 

10. Driotura gammaroides (Van Duzee). 

11. Euscelis exitiosus (Uhler). 

12. Euscelis striolus (Fallen). 

13. Eiiscelis comma (Van Duzee). 

14. Euscelis bicolor (Van Duzee). 

(270) 



PLATE XXI II. 




««*fes^ 








10 




(271) 



PLATE XXIV. 

1. Entettix cinctus Osborn and Ball. 

2. Entettix strobi (Fitch). 

3. Phlepsius spatulatus Van Duzee. 

4. Phlepsius excultus (Uhler). 

5. Phlepsius irroratus (Say). 

6. Acinopterus acuminatus Van Duzee. 

7. Chlorotettix spatulatus Osboin and Ball. 

8. Chlurotettix galhanatus Van Duzee. 

9. Cicadula punctijrons var. rcpleta Fieber, 

10. Balclutha punctata (Thunberg). 

11. Balclutha impicta (Van Duzee). 

12. Eugnathodus abdominalis (Van Duzee). 

13. Dikraneura abnormis (Walsh). 

14. Empoasca obttisa Walsh. 

(272) 



PLATE XXIV. 





3 




8 





(273) 



PLATE XXV. 

1. AijaUiopsis novella (Say). 

2. Aceratagallia uhleri \i\nT>uzee. 

3. hUocerns snowi Gillette and Baker. 

4. Idiocerus ramentosus (Uliler). 

5. Idiocerus pallidus Fitch. 

6. Idiocerus duzeei Provanchei'. 

7. Idiocerus verticis (Say). 

8. Idiocerus scurra (Gerniar). 

9. Idiocerus nervatus Van Duzee. 

10. BythoHcopus apicalis (Osborn and Ball). 

11. Bythoscopus iniscUus (Stal). 

(274) 



Readio: Ovipositors of Cicadellid.e. 



27.") 



I'LATE XXV 




276 The University Science Bulletin. 



PLATE XXVL 

1. Macropsis viiidis (Fitch). 

2. Macropis suturalis (Osborn and Ball). 

3. Oncopsis distinctus (VanDuzee). 

4. Oncometopia unda'a (Fabricius). 

5. Oncometopia lateralif^ (Fabricius). 

6. Homalodisca triquelra (Fabricius). 

7. Aulacizes irrorata (Fabricius). 

8. Cicadella circellata (Baker). 

9. Cicadella hieroglyphica (Say). 



Readio: Ovipositors of Cicadellid^. 



277 



PLATE XXVI. 




278 The University Science Bulletin. 



PLATE XXVII. 

1. DrcEculacephala noveboracensis (Fitch). 

2. Drceculacephala mollipes (Say). 

3. Drceculacephala reticulata (Signoret). 

4. Errhomenellus montaniis Baker. 

5. Helochara communis Fitch. 

6. Pagaronia tripunctata (Fitch). 



Rkadio: Ovipositors of CicAnKLLin.i': 



279 



PLATE XXVIT. 




280 The Ut.'iversity Science Bulletin. 



PLATE XXVIII. 

1. Kolla hartii (Ball). 

2. Kolla geometrica (Signoret). 

3. Kolla bifida (Say). 

4. Graphocephala coccinea (Forster), 

5. Gypona bimaculata Spangberg. 

6. Gypona angulata Spangberg. 

7. Gypona pectoralis Spangberg. 

8. Gypona octo-lineata (Say). 



Readio: Ovipositors of Cicadellid.e. 



281 



PLATE XXVIII. 




282 The University Science Bulletin. 



PLATE XXIX. 

1. Xe^torephahis: puUcariun Van Duzce. 

2. Xerophloea viridis (Fabricius). 

3. Stroggylovcphalu.s agrcstis (Fallen). 

4. Dorycephalus platyrhynchns Osborn. 

5. Mcmnonia consobrina Ball. 

6. Spangbergiella mexicana Baker. 

7. Parabolocratus fiavidus Signoret. 

8. Hcculus lineadus (Uhler). 

9. Meaamia straim)ua (Osborn). 

10. Mesamia viiellina (Fitch). 

11. Aligia jiicunda (Uhler). 



Rkadio: ()\'ii'081toks of ('i('Ai)KLi>in.i':. 



283 



PLATE XXIX. 




284 The University Science Bulletin. 



PLATE XXX. 

1. Scaphoideiis scalaris Van Duzee. 

2. Scaphoideus immistus (Say). 

3. Platymetopius acutus (SajO- 

4. Plalymetopius cinereus Osborn and Ball. 

5. Platymetopius frontalis Van Duzee. 

6. Deltocephalus reflcxus Osborn and Ball. 

7. Deltocephalus weedi Van Duzee. 

8. Deltocephalus inimicibs (Say). 

9. Deltocephalus flavicosta Stal. 

10. Deltocephalus debilis Uhler. 

11. Deltocephahis parvulus Gillette. 

12. Deltocephalus collinus Boheman. 



Readio: Ovipositors of CiCADELLiDyE. 



285 



PLATE XXX. 




286 The University S(;ience Bulletin. 



PLATE XXX r. 

1. Euscelis exitiosus (Uhler). 

2. Euscelis striolus (Fallen). 

3. Euscelis anthracinus (Van Dnzf c) 

4. Eusceliji comma (Van Diizee). 

5. Euscelis curLesii (Fitch). 

6. Euscelis bicolor (Van Duzee). 

7. Eutettix strobi (Fitch). 

8. Eutettix cinctus Osborn and Ball. 

9. Aconura argenteolus (Uhler). 

10. Nephotettix curtipennm (Gillette and Baker). 

11. Driotura garnmaroides (Van Duzee). 



Readio: Ovipositors of Cicadellid.e. 



287 



PLATE XXXI. 




288 The University Science Bulletin. 



PLATE XXXII. 

1. Acinopterus viridis Ball. 

2. Acinopterus angulatus Lawson. 

3. Acinopterus acuminatus Van Duzee. 

4. Thamnotettix longulus Gillette and Baker. 

5. Thamnotettix clitellarius (Say). 

6. L'hlorotettix spatulatus Osborn and Ball. 

7. Chlorotettix galbanatv^s Van Duzee. 

8. Phlepsiu^ irroratus (Say). 

9. Phlepsdus excultuji (Uhler). 

10. Phlepsius spatulatus Van Duzee. 



Readio: Ovipositors of Cicadellid^. 



289 



PIATE XXXII. 




290 The University Science Bulletin. 



PLATE XXXIII. 

1. Tinohregmus vittatus Xnn Diizee. 

2. Jassus olitorius Say. 

3. Cicadula ■punctifrons var. repleta Fieber. 

4. Balclutha punctata (Thunberg). 

5. Balclutha impicta (VanDuzee). 

6. Eugnathodus abdominalis (Van Diizee). 

7. Dikrancura ahnormis (Walsh). 

8. Erythroneura tricincta Fitch. 

9. Erythroneura vulnerata Fitch. 

10. Empoasca trifasciata Gillette. 

11. Empoasca smaragdula (Fallen). 

12. Empoasca obtusa Walsh. 

13. Empoasca mali (Le Baron). 

14. Empoasca livingstoni Gilli tie. 



Readio: Ovipositors of Cicadellid.k. 



291 



PLATE XXXIII. 




INDEX. 

PAOK 

Abdomen of the female, The 218 

abdominalis, Eugiuithodus 260 

abnormis, Dikraneura 261 

Aceratagallia 

uhleri 224 

Acinopterus 255 

acuminatus 255 

angulatus 256 

viridis 255 

Aconura 

argentiolus 248 

Acucephalini 238 

acuminatus, Acinopterus 255 

acutus, Platymetopius 243 

Agalliopsis 224 

novella 224 

agrestis, Stroggj-locephalus 238 

Aligia 

jucunda 240 

angulata, Gypona 237 

angulatus, Acinopterus 256 

anthracinus, Euscelis 250 

apicalis, Bythoscopus 228 

argentiolus, Aconura 248 

Aulacizes 

irrorata 230 

Balclutha 260 

impicta 260 

punctata 260 

bicolor, Euscelis 252 

bifida, Kolla 232 

bimaculata, Gypona 237 

Bythoscopinse ; 224 

Bythoscopus 228 

apicalis 228 

miscellus 229 

Chlorotettix 257 

galbanatus 258 

spatulatus 258 

Cicadella 231 

circellata 231 

hieroglyphica 231 

Cicadellinae 229 

Cicadidaj 217 

Cicadula 

punctifrons var. repleta 259 

cinctus, Eutettix 252 

(293) 



294 The University Science Bulletin. 

PAGE 

cinereus, Platymetopius 244 

circellata, Cicadella 231 

clitellarius, Thamnotettix 256 

coccinea, Graphocephala 234 

collinus, Deltorephalus 247 

comma, Euscelis 251 

communis, Helochara 233 

Conclusions 264 

consobrina, Memnonia 238 

curtesii, Euscelis 251 

curtipennis, Nephotettix 248 

dehilis, Deltocephalus 247 

Deltocephalus 245 

collinus '. 247 

debilis 247 

flavacosta 246 

inimicus 246 

parvulus 247 

reflexus 245 

weedi 245 

Dikraneura 

abnormis 261 

distinctus, Oncopsis. 228 

Dorycephalus 

platyrhynchus 239 

Dripculacephala 234 

moUipes 234 

noveboracensis 235 

reticulata 235 

Driotura 

gammaroides 249 

duzeei, Idiocerus 225 

Empoasca 261 

livingstoni 262 

roali 263 

obtusa 262 

smaragdula 262 

trif asciata 26 1 

Errhomenellus 

montanus 236 

Erythroneura 263 

tricincta 263 

vulnerata 264 

Eugnathodus 

abdominalis 260 

Euscelis 249 

anthracinus 250 

bicolor 252 



Readio: Ovipositors of Cicadellid.^*:. 295 

rA(i !■; 

comma 251 

curtosii 251 

cvitiosiis 250 

strioliis 250 

Eutettix 252 

cinctus 252 

sirol)! ■ 253 

exitiosus, iMiscelis 250 

cxcultus. I'hIoi)sius 254 

flavacosta, Deltooephaius 246 

flavidus, Parabolocratus 240 

frontalis, Platymetopius 244 

galbanatus, Chlorotettix 258 

gammaroides, Driotura 249 

geometrica, Kolla 232 

Graphocephala 

coccinea 234 

Uypona 236 

angulata 237 

biraaculata 237 

ooto-lineata 237 

pectoralis 237 

Gyponinae 236 

hartii, Kolla 233 

Hecalus 

lincatus 239 

Helochara 

communis 233 

hieroglyphica, Cicadella 231 

Homalodisca 

triquetra 230 

Idiocerus 224 

duzeei 225 

iiervatus 225 

pallidas 225 

ramentosus 226 

.snowi 226 

scurra 226 

verticis 226 

imniistus, Scaphoideus 243 

impicta. Balclutha 260 

inimicus, Deltocephalus 246 

Introduction 217 

irrorata, Aulacizes 230 

irroratus, Phlepsius 254 

Jassinae 238 

Jassini 239 



296 The University Science Bulletin. 

Jassus PAGE 

olitorius 258 

jucunda, Aligia 240 

Kolla 232 

bifida 232 

geometrica 232 

hartii 233 

lateralis, Oncometopia 230 

lineatus, Hecaliis 239 

livingstoni, Empoasca 262 

longulus, Thamnotettix 257 

Macropsis 227 

suturalis 227 

viridis 227 

mali, Empoasca 263 

Memnonia 

consobrina 238 

Mesamia 241 

straminea 241 

vitellina 241 

Methods 223 

mexicana, Spangbergiella 240 

miscellus, Bythoscopus 229 

mollipes, Drseculacephala 234 

montanus, Errhomenellus 236 

Nephotettix 

curtipennis 248 

nervatus, Idiocerus 225 

noveboracensis, Drseculacephala 235 

novella, Agalliopsis 224 

obtusa, Empoasca 262 

octo-lineata, Gypona 237 

olitorius, Jassus 258 

Oncometopia 229 

lateralis 230 

undata 229 

Oncopsis 

distinctus ' 228 

Oviposition .' 220 

Ovipositor 

Description of 220 

Taxonomic use of 222 

Pagaronia 

tripunctata 236 

pallidus, Idiocerus 225 

Parabolocratus 

flavidus 240 

parvulus, Deltocephalus 247 

pectoralis, Gypona 237 






Readio: Ovipositors of Cicadellid.^:. 297 

PAOK 

Phlepsius 253 

cxcultus 254 

irrorat us 254 

spatulatus ■. . . 253 

Photography of ovipositors 218 

Platyinotoi)ius 243 

aculus 243 

cinereus 244 

frontalis 244 

platyrhyiu'hus, Dorycephalus 239 

pulicarius, Xestocephalus 239 

punctata, Balclutha 260 

punctifrons ver. repleta, Cicadula 259 

pygofer 220 

ramentosus, Idiocerus 226 

reflexus, Deltocephakjs 245 

repleta, var., Cicadula punctifrons 259 

reticulata, Dra'culacephalu 235 

scalaris, Scaphoideus 242 

Scaphoideus 242 

immistus 243 

scalaris 242 

scurra, Idiocerus 226 

Segments, Number of 218 

smaragdula, Empoasca 262 

snowi, Idiocerus 226 

Spangbergiella 

mexicana 240 

spatulatus, Chlorotettix 258 

spatulatus, Phlepsius ' 253 

Spiracles, Number of 219 

straminea, Mesamia 241 

striolus, Euscelis 250 

strobi, Eutettix 253 

Stroggylocephalus 

agrestis 238 

suturalis, Macropsis 227 

Thamnotettix ■ 256 

clitellarius 256 

longulus 257 

Tinobregmus 

vittatus 259 

tricincta, Erythroneura 263 

trifasciata, Empoasca 261 

triquetra, Homalodisca 230 

tripunctata, Pagaronia 236 

Tj'phlocybini 201 

uhleri, Aceratagallia 224 

undata, Oncometopia 229 



298 The University Science Bulletin. 

PAGE 

vertiL-is, Idiocerus 226 

viridis, Acinopterus 255 

viridis, Macropsis 227 

viridis, Xerophloea 238 

vitellina, Mesamia 241 

vittatus, Tinobregmus 259 

vulnerata, Erythroneura 264 

weedi, Deltocephalus 245 

Xerophloea 

viridis 238 

Xestocephalus 

puliearius 239 



I 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 9— October, 1922. 

(Whole Series, Vol. XXIV, No. 9.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

Life History Notes on Two Species of Saldid.e (Heteroptera), 

Grace Olive Wiley. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE, KAN. 



Entered at tlie post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 9. 



Life History Notes on Two Species of Saldidse 
(Hemiptera) Found in Kansas.* 

BY GRACE OLIVE WILEY. 

THE family characteristics of the saldids are so well known that 
a description need not be given here. A few brief notes on 
certain Kansas species, however, may be of interest. 

Nymphs in the third and fourth instars were taken June 1, 1920, 
along a little ravine leading to a pond or pool which contained w^ater 
only during the spring and early summer when rains were frequent. 
This little freshet and pond were in a small pasture just outside the 
city limits of Chanute, Kan. Here, within a few blocks of home, I 
found two different species of saldids, though in not very plentiful 
numbers. 

Knowing that no Kansas saldid had yet been reared, I captured 
all the lively little fellows I could and sat about the task of rearing 
them. One was a large black species, the other a much smaller 
species. 

THE LARGE BLACK SALDID. 
Saldula major (Pro\\). 

June 12 there were two adults, and by June 18 the others had 
reached the adult state and were mating. June 21 I found twelve 
newly laid eggs on a blade of grass. These hatched July 3. With 
what pleasure and interest these active little fellows were watched! 
The adults were shy, but very inquisitive, and when food was placed 
in their dish they were always ready to run up and take a look at it, 
curiosity-bent, it seemed; or when removing food that had been 
given to them the day previous, they were equally curious to see 
what was going on. Not so with the smaller species; they were 
always trying to get away or hide. 

*I wish to thank Dr. H. B. Hungerford for his help and kindly criticism and for the loan 
of a binocular from the University. 

(301) 



302 The University Science Bulletin. 

The nymphs liked to stay hidden most of the time, but would 
come out and feed quite readily. It was interesting to see one of 
the little fellows prodding around an insect for a soft spot in which 
to insert its needlelike beak. Nor does it disturb one of them in the 
least to remove the insect and place same under the binocular to 
watch the process of feeding. 

The newly emerged nymphs, either from the egg or molt, are very 
bright red in color, with eyes, antennae, beak and legs very dark. 
They become dark, however, in a very short time, except the nymphs 
which emerge from the eggs; these are red for more than a day. 
The adults are very pale pink or yellowish when they first emerge, 
with eyes, antennae, beak and legs very dark. They become dull 
black in a short time, with rather pale, obscure markings on the 
wing covers. 

DESCRIPTION OF STAGES. 

See plate XXXIV. 
THE EGG. 

Size. Length about 1.2 mm.; width across mdest part, .3 mm. 

Color. Pearly white and transparent when first laid, becoming yellow in 
color three days later. The fourth-day egg shows red eyelike spot, and two 
days later the entire egg becomes red in color; another day later, making seven 
daj's from time egg was laid, it is blood red with verj' dark, eyelike spots. 
When nine days old the egg is orange red, and on the twelfth day the nymph 
emerges. A few hours before hatching the egg becomes pale or whitish in 
color. 

Shape. General shape shown by drawing. It is elongate, larger and more 
broadly rounded at one end, tapering and much smaller at the other end, with 
dorsal part arched. 

FIRST INSTAR. 

Size. Total length of one day-old nymph, about 1.1 mm.; width across 
abdomen, .6 mm. 

Color. Red to light reddish brown, with dark spot on fifth abdominal seg- 
ment, on the median line of the dorsal part. Eyes, legs, beak and antennae 
black. 

Structural peculiarities. Rostrum reaching to the apex of the posterior coxae. 
Antennae four-segmented, last joint darker in color and larger than the others. 
Tai-si one-jointed. Length of first stage, four days. 

SECOND INSTAR. 

Size. Length about 1.8 mm.; width across abdomen about 1 mm. 

Color. Dark reddish brown on dorsal part,^with ventral part red. Venter 
very dark brown, nearly black. 

Structural peculiarities. Antennae four-segmented, with last segment larger 
than the other segments. Rostrum reaching to the base of the posterior 
coxae. Antennae and legs covered with very fine black hairs, some longer and 
more sparsely placed than the others. 

Shape. A trifle stouter than first-instar nymph. 



Wiley: Saldid.e. 303 

THUJD IN STAR. 

Size. Lenjrth from 2.25 to 2.5 inin,; width across abdomen from 1.3 mm. to 

1.4 nun. 

Color. Color on the <lorsum very dark brownish black, with ventral parts 
brown. 

Shape. Much the same as the second instar. only much sto\iter. 

FOIKTH INST.\R. 

Size. Length. 3.25 mm. to 3.5 mm.; width across abdomen, 1.9 mm. to 2 mm. 

Color. Shining black o\er entire part ; most of the ventral part black ex- 
cept the throat and around the legs, which is slightly paler in color, also the 
anterior and lateral jiarts of the carina or keel of the abdomen, which is 
whitish in color; venter very dark. 

Shape. Decidedly stouter than third instar and wing iiads now much in 
evidence. 

Structural peculiarities. Rostrum reaching to the apex of the posterior 
coxae. Ventral part of the abdomen carinated. Antenna^ long, rather stout, 
and covered, as in the third instar. with fine black hairs. 

FIFTH IXST.\R. 

Size. Length from about 4.5 mm. to 4.8 mm.; width across abdomen at 
widest part, 2.5 mm. to 2.625 mm. 

Color. All of dorsum shining black in color. Ventral parts same as in 
fourth instar, as are also the antennae and legs. Second joint of antennae de- 
cidedly longer and more slender than any of the other joints. Rostrum reach- 
ing to the base of the posterior coxiv. Eyes emarginate. Wing pads reaching 
to the apex of the second abdominal segment. 

THE .ADULT. 

Size. Female: length of the body, not including the antennae and legs, 

6.5 mm. to 6.875 mm.; width, 2.8 to 3 mm. Male: length, 6.25 mm.; width, 
2.625 to 2 mm. 

Color. Entire insect dull black with a few obscure, pale markings. Body 
oblong, oval. Ocelli slightly apart. Head long and tapering to the front. 
Eyes large, emarginate and strongly exserted. Scutellum rather flat with a de- 
pression in center of disk. The posterior lateral margins of pronotum and the 
anterior margins of hemelytra slightly refiexed. Rostrum long and slender, 
first joint short and thick, second joint very long, the tip reaching to the base 
of the middle coxae; third joint reaching to the base of the posterior coxae. 
Color of beak black and shining, with tip slightly paler. Antennae very black, 
stout and long; basal joint stout and considerably shorter than third or fourth 
joints; second joint very long and slender; last two joints nearly the same 
length; last joint slightly longer than the next to last. Body covered with 
sparse, closeh' appressed, golden pubescence. Membrane furnished with four 
areas. 

Oviposition. The eggs are deposited at the base of grass blades, 
or are thrust \vith the sharp ovipositor through the bhndes of grass. 
(See plate XXXIV. j 



304 



The University Science Bulletin. 



Incubation. Twelve days. 

Maturity. Adults come from eggs in twenty-eight days. 

General notes. Length of first instar, four days; of second in- 
star, three days; of third instar, three days; of fourth instar, two 
or three days; of fifth instar, four days. No ocelli are present in any 
of the nymphal stages. 

MEASUREMENTS IN MILLIMETERS OF NYMPHS AND ADULTS OF LARGS SPBCIE=i 

KANSAS SALDID. 





3 
» 


Body measurements, m m. 


Leg measurements, m ra. 




? 


3 
TO 




>- 
O 

CO 


_3- 

w 

3- 

g 


Length, antenna . . 


CO 

$ 


Fore leg. 


Middle leg. 


Hind leg. 






3- 










3 


a; 
so' 


H 
2. 


3 

-1 


St 
5' 


3_ 


Femur 




-• 


1st 


1.1 


.6 


.4 


.5 


.6 


.7 


.3 


.25 


.1 


.35 


.3 


.15 


.4 


4 


.2 


2d 


1.8 


1.0 


.7 


.75 


1.0 


1.1 


.5 


.4 


.2 


.5 


.45 


.25 


.6 


.7 


.3 


3d 


2,5 


1.4 


.9 


1.0 


1.6 


1.6 


.6 


,5 


.25 


.7 


.6 


.3 

.4 


.8 


1.0 


.4 


4th 


3.5 


1.9 


1.1 


1.3 


2.0 


2,0 


.9 


.7 


.3 


1.0 


1.0 


1.2 


1.5 


.6 


5th 


4.8 


2.5 


1.4 


1.9 


2.7 


2.7 


1.2 


1.0 


.5 


1.5 


1.2 


.6 


1.6 


2.2 


.9 


6th 9 


6.5* 6.75t 


2.8 


1.5 


2.0 


3.5 


3.0 


1.5 


1.3 


.7 


1.6 


1.6 


.8 


2.0 


3.0 


1.25 


6th 9 


6.0* 7.0t 


3.0 


1.5 


2.25 


3.5 


3.5 


1.8 


1.4 


.7 


1.8 


1,6 


.7 


2.2 


3.0 


1.25 


6th c? 


6.0 


2.8 


1.5 


2.0 


3.5 


3.0 


1.5 


1,4 


.7 


18 


16 


.8 1 2.0 


3.0 


1.25 



* Venter. t Elytra. 

These measurements were made with micrometer, Bausch and Loml) binocular. 6.1 oculars, 55 mm. objec- 
tives. By placing millimeter ruler under binocular, micrometer measured ten lines to the millimeter. Thus: 10 
lines micrometer = 1 mm. 

SMALL SPECIES OF KANSAS SALDID. 

Saldula pOlUpes Fabr. (?)* 

From adults reared from nymphs captured June 1, 1920, along 
with nymphs of a large black species of saldid, I obtained, June 
23, several clear creamy- white eggs, thrust in the stems and blades 
of grass growing in the jars in which these saldids were confined. 
These were not the first eggs laid, however, as I also found several 
small nymphs, but the eggs had been so cleverly hidden that I had 
not found them. These eggs hatched July 1 to 3, and became adults 
sixteen days later. Eggs from this second generation hatched 
August 13 and August 20, becoming adults also in from sixteen to 
seventeen days from the time the nymphs emerged from the eggs. 

On the 28th of July I went to Texas to join my husband, who was 
in the employ of an eastern oil company. I could not think of 



*Mr. Hung.-rforcl says this .American species is not the same as the Europtan species. 



Wiley: 8aldid^. 305 

giving up tlic life histories i)f the two species of saldids I was rearing, 
so phicing all the little glasses containing them in a covered basket, 
I took them with me. No doubt the people on the train thought I 
was taking a lunch with me, or perhaps a pet dog or cat! I wonder 
how many ladies would have slept well that night had they known 
that the basket contained live bugs! 

In all the rearings I fed dead flies and other soft-bodied insects,, 
chiefly mirids (capsids) and cicadellids (jassids), as these were 
usually easier to obtain in large numbers either by sweeping or at 
the light at night. 

DESCRIPTION OF STAGES. 

See Plate XXXV. 
THE EGG. 

Size. Length about .6 mm.; width, .15 mm. 

Color. Clear, creamy-white when tirst laid, changing to yellow. Duration 
of egg stage. 8 to 10 days. 

Shape. Elongate-cylindrical; one end broadly rounded and considerably 
larger than the other end. Dorsal jiart arched. 

FIRST INSTAR. 

Size. Length of body about .8 mm.; width of abdomen at widest part, 
.4 mm. 

Color. Greenish brown with dark spot on the dorsum of abdomen along 
median line. 

Structural peculiarities. Mostly head, with large, reddish eyes; four-seg- 
mented slender antennae, fourth segment stouter than the others. Duration of 
first instar, four days. 

SECOND INSTAR. 

Size. Length, 1. mm.; width, .6 mm. 

Color. Light green with eyes and antennse dark reddish brown. Dark spot 
on the dorsum of abdomen. 

Shape. Much the same as first instar only stouter. Duration of instar, 
three days. 

THIRD INST.AR. 

Size. Length, 1.5 mm.; width, .8 mm. 

Color. Head and thorax yellowish; third and fourth abdominal segments 
dark green with spot on the dorsum orange yellow; remainder of abdomen 
yellowish green. Eyes and tip of antennae dark brown. 

Structural peculiarities. Tarsi segmented as in all the other instars; rostrum 
reaching to the posterior coxae. Duration of third instar, three days. 

FOLTRTH INSTAR. 

Size. Length of body, 2 mm.; width of abdomen, 1.2 mm. 

Color. Grayish flecked with brown. 

Shape. Nymph much stouter than in the third instar. and wing pads reach- 
ing to the base of the second abdominal segment. Beak reaching to the apex 
of the posterior coxae. Duration of fourth instar, 3 days. 



306 The University Science Bulletin. 

FIFTH INSTAR. 

Size. Total length, 2.7 mm. to 3 mm.; width, 1.5 mm. to 1.6 mm. in wides^t 
portion. 

Color. Grayish brown speckled or mottled. 

Structural peculiarities. Antennae slender, fovir-segmented as in the other 
instars; fourth segment larger and stouter than the others; second segment 
longest and very slender; body, legs and antennae covered with spai-se, short, 
brown pubescence. Rostrum reaching to the apex of the posterior coxae. 
Wing pads reaching to base of third abdominal segment. Duration of fifth 
instar, three days. 

THE ADULTS. 

Size. Length of the entire body, not including antennae and legs: Female: 
3.5 mm. to tip of abdomen; 3,9 to 4 mm. to tip of hemelytra; width of abdo- 
men across the widest part, 1.6 mm. to 1.7 mm. Male: length 3 mm. to tip 
of hemelytra; width, 1.5 mm. 

Description. Oblong-ovate, black above, closely invested with minute yellow 
pubescence; eyes large, rather oblong, brown, and very prominent. Ocelli 
reddish and slightly apart. Clypeus and tylus straw yellow; rostrum black 
and reaching upon the posterior coxae. Ventral part body jet black; sternum 
and pectus closely appressed with silvery-white pubescence. Legs pale with 
dark markings. Pronotum and scutellum rather flat, disk of pronotum slightly 
elevated and with a minute depression in center of disk. Scutellum a Uttle 
longer than broad. Hemelytra brownish black with grayish-white markings. 
Clavus with oblong spot at the apex. Corium with two squarish spots near 
the upper, outer margin and two very small spots on the posterior margin near 
membrane. Embolium mostly grayish white, there usually being three dark 
spots along the median nerve connected by a dark-colored line or nerve. 
Membrane gray with dark veins and furnished with four areas, each having 
one or more smoky spots therein. Antennae slender, four-segmented; basal 
segment stoutest, dark underneath, pale above; second segment very slender 
and almost twice the length of the first segment; third and fourth segments 
about equal in length and stouter than the second. 



Wiley: Saldid.^. 



307 



ME.\SUREMENTS IN MILLIMETERS OF NYMPHS AND ADULTS OF SMALL SPECIES 

KANSAS SALDID. 







Body measurements, mm. 


Leg measurements, mm. 




1 


5* 


31 


>• 

1 


a! 

at 

cr 
o 

c 


a- 
a 
o 




Fore leg. 


Middle leg. 


Hind leg. 






D- 










a 

c 


6= 


3. 


3 
c 
-1 


s: 

CO 


i. 


■4 

B 
c 




I 


1st 


.8 


.4 


.350 


.3 


A 


.4 


.125 


.12 


.065 


.15 


.125 


.070 


.16 


.25 


.080 


2d 


1.0 to 
1.1 


.6 


.4 
.5 


.45 
.525 


.5 


.5 
.6 


.2 


,2 


.1 


.3 


.2 


.125 


.3 


.4 


.175 


3d 


1.5 


.8 


.6 


.7 


.8 


.9 


.4 


.35 


.1 


.4 


.35 


.15 


.5 


.6 


.2 


4th 


1.8 to 
2.1 


1. 
1.2 


.6 

.7 


.8 
.9 


1.0 


1.1 

1.2 


.4 
.5 


.4 


.2 


.5 

.6 


.4 
.5 


.2 


.6 

.7 


.7 
.8 


.25 
.3 


5th 


2.7-2.9 
to 3 


1.5 
1.6 


.9 
1.0 


1.25 
1.3 


1.5 


1.5 
1.6 


.7 


.6 


.25 


.8 
.9 


.7 


.3 


.9 
1.0 


1.1 


.5 


6th 9 


3.5* 4.0t 
3.9 


1.6 
1.7 


1.0 


1.3 


1.6 
1.7 


1.6 
1.7 


.9 


.7 


.3 


.9 


.8 


.35 

.4 


1.1 


1.6 


.6 


6thc? 


3.0 to 

elj'tra 


1.5 


.9 


1.2 


1.5 


1.5 


.7 


.6 


.25 


.8 


.7 


.3 


.9 


1.5 


.5 



* Venter. t Elytra. 

Measurements made with micrometer same as in large species, 10 lines micrometer = 1 millimeter. 



308 The University Science Bulletin. 



PLATE XXXIV. 
THE LARGE SALDID. 

Saldula major (Prov.). 

Fig. 1. Stem of grass with some of the blades removed, showing egg of 
saldid. 

Fig. 2. Under side of grass blade, showing tips of eggs through the opening 
made by the sharp ovipositor. 

Fig. 3. Upper side of the same blade shown in figure 2, showing the eggs 
and how they were thrust through the leaf. 

Fig. 4. Egg in a grass blade and part of the blade torn away to show the 
tip of the egg on under surface. 

Fig. 5. The egg. 

Fig. 6. Nymph just out of egg, still in postnatal molt. 

Fig. 7. First-instar nymph. 

Fig. 8. Second-instar nymph. "n 

Fig. 9. Fourth-instar nymph. 

Fig. 10. Third-instar nymph. 

Fig. 11. Fifth-instar njnnph. 

Fig. 12. Adult male. 

Fig. 13. Adult female, ventral view. 

Fig. 14. Adult female, dorsal view. 



Wiley : Saldid.*:. 



309 



PLATE XXXIV 




310 The University Science Bulletin. 



PLATE XXXV. 
THE SMALL SALDID. 

Saldula pi/liipes Fabr. (?) 

Fig. 1. The egg. 

Fig. 2. First-instar nymph. 

Fig. 3. Second-instar nymph. 

Fig. 4. Third-instar nymph. 

Fig. 5. Fourth-instar nymph. 

Fig. 6. Fifth-instar nymph. 

Fig. 7. Adult female. 

Fig. 8. Adult male. 



D 




Wiley: Saldid.e. 
PLATE XXXV. 



311 






5. 






d 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 10— October, 1922. 

(Whole Series, Vol. XXIV, No. 10.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

A Problem in the Relation of Temperature to Rate of In- 
sect Development P. A. Glenn. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE, KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 10. 



A Problem in the Relation of Temperature to 
Rate of Insect Development. 

By p. A. GLENN, 

Chief Inspector, Division of Plant Industry, Illinois State Department of 
Agriculture, Urbana, 111. 

EVEN the most casual observer is familiar with the fact that low 
temperatures inhibit growth in plants and animals, that hiber- 
nation or death will be evidenced by temperatures approaching 
freezing, and that high temperatures favor rapid development. It 
is only in late years that biologists have attempted to ascertain 
with accuracy the reaction of various plants and animals to differ- 
ent degrees of heat. It has now been quite definitely demonstrated 
that development will take place only when the temperature is 
above a certain point, called the zero of development, or the 
threshold of development; that within a certain range of tempera- 
tures above this point the increase in the rate of development is 
approximately proportional to the rise in temperature; and that 
under given conditions of high moisture, evaporation and other 
physical environmental factors aside from heat, there is a tempera- 
ture constant for each period in development, which is equal to the 
product of the period by the average temperature above the thres- 
hold of development. 

This range of temperatures within which the rate of development 
increases as the temperature rises is bounded at the lower end by 
the threshold of development, and at the upper end by what I 
shall call the degree of the maximum rate of development. It is 
found that at temperatures near the lower limit of this range the 
rate of development varies somewhat faster than the rate of change 
in temperature, and at temperatures near the upper limit the rate 
of development varies somewhat more slowly than the rate of change 

(317) 



318 



The University Science Bulletin. 



of temperatures. These facts are of importance to scientists who 
wish to ascertain the exact relations between temperature and 
development, but for practical purposes these slight variations 
need not be taken into consideration, especially when the varying 
out-of-door temperatures are to be used as a basis of study. 

The problem, therefore, dealt with in this paper may be stated 
as follows: Given the length in days of the period of development 
at different average daily temperatures, and the average daily 
temperatures for each of the periods, to find the threshold of de- 
velopment, the degree of the maximum rate of development, the 
temperature constant, the effects of temperatures above the de- 
gree of the maximum rate of development, and how to make cor- 
rections in the temperature factor when the temperature for a part 
of the time during the period was above the degree of the maximum 
rate of development. 

The problem is a simple one when constant temperatures are 
used, but with widely varying daily temperatures as a base for 
study the problem is more difficult, since the average daily temper- 
atures for the periods must be used and in nearly all observations 
these are influenced by temperatures below the threshold of develop- 
ment or temperatures above the degree of the maximum rate of 
development. 

In this paper the method of procedure will be illustrated by studies 
on the pupal period of the Carpocapsa pomonella, based on observa- 
tions out of doors on the pupal period of 3,817 pupae at mean daily 
temperatures varying from 52.6° F. to 82.6° F. 

The following table gives the complete results of the study : 



RELATION OF TEMPERATURE TO LENGTH OF PUPAL PERIOD. 


Temper- 
ature 
range. 


Generation. 


Number 

of 
obser- 
vations. 


Har- 
monic 
average 

of 
period. 


1 
P 


Aver- 
age 
mean 
daily 
temper- 
ature. 


Average daily 
day-degrees. 


Total day- 
degrees. 


52-t- 


2(87-F) 


(52-F)- 
2(87-F) 


52+ 


(52+)- 
2(87+) 


52-55 


Hib 


2 
348 
976 
243 
218 
175 
221 
247 
517 
481 
133 
256 


45.5 
35.2 
34.0 

27.7 

13.8 

12.7 

11.5 

10.7 

10.0 

9.4 

9.4 

9.2 


.021 
.028 
.029 
.036 
.072 
.078 
.086 
.093 
.099 
.105 
.106 
.108 


52.6 
55.7 
56.1 
58.5 
69.2 
70.7 
73.1 
74.9 
76.7 
79.1 
80.8 
82.6 


5.1 

6.8 

7.0 

8.6 

17.3 

18.7 

21.1 

22.9 

24.7 

27.1 

28.8 

30.6 


";64' 

.24 

.42 

.88 

1.63 

3.52 

4.12 


5.1 

6.8 

7.0 

8.6 
17.3 
18.7- 
20.8 
22.5 
23.8 
25.5 
25.3 
26.5 


236 
237 
240 
241 
238 
238 
242 
245 
248 
256 
272 
283 


236 


54-55 


Hib 


237 


56-57 


Hib 


240 


58-59 . . . 


Hib 


241 


68-69.... 
70-71.... 
72-73.... 
74-75.... 
76-77. . . . 
78-79 


First and second. 
First and second. 
First and second. 
First and second, 
First and second, 
First 


238 
238 
240 
241 
239 
240 


80-81 


First 


239 


82-83 


First 


245 








52-83.... 




3,817 


14.4 


.669 




17.3 


.77 


16.6 


251 


240 



Glenn: Insect Development. 319 

Tlio usual method for determining the threshold of development 
is to use the reciprocals of the periods and the average mean daily 
temperatures as the coordinates and plot the one against the other, 
then draw a line which best fits the points, and the point where it 
crosses the temperature axis is the threshold of development. This 
method serves quite well when the temperatures are constant, but 
the more widely the temperature varies during the daily variations 
the less accurate are the results. The accompanying figure is really ■ 
four figures placed on the same sheet, and represents the points 
in the various positions assumed by them at the different stages 
in the solution of the problem. 

The circles represent the position of points resulting from plotting 
the reciprocals of the periods against the average mean daily 
temperatures. These points do not fall in a straight line, but at the 
lower end they are too far to the left, because the average mean 
daily temperatures are the averages of temperature readings below 
the threshold of development (ineffective temperatures) as well as 
readings above this point, and the points at the upper end are too far 
to the right, because the average mean daily temperatures here are 
averages of readings above the degree of the maximum rate of 
development (retarding temperatures) as well as readings below 
this point. 

A line that would fit these points would be too flat. Now the 
points that are least affected by these ineffective and these retard- 
ing temperatures are those between the average mean daily tempera- 
tures of 68 and 72. A line drawn through these two points cuts 
the temperature axis at 51.92 degrees. This is so nearly 52 de- 
gress that, to avoid fractions, 52 was assumed to be the approximate 
threshold of development. 

The average daily degrees above 52 were then computed with 
results as shown in column 7. The points marked by dots were 
plotted by using the reciprocals of the periods and the average 
daily temperatures above 52 degrees. It will be seen that at the 
lower end of the line the points come into line with the two points 
used to establish the line, but at the upper end of the figure they 
are in the same position as the points located by plotting reciprocals 
against average mean daily temperatures. 

It was necessary now to ascertain the degree of the maximum rate 
of development. The only way to determine this point is by trial. 

It was assumed at first that temperatures above the degree of the 
maximum rate of development were ineffective and did not ac- 



320 



The University Science Bulletin. 



'^ 



--4 •' 



^4 ,^ 







Glenn: Insect Development. 321 

celerate the rate of development. On this supposition 84, 85, 86, 
87, 88 and 89 were taken in turn as the degree of the maximum rate 
of development and the proper corrections made. This did not bring 
the points back into line, as the points higher up still stood too far 
to the right. It was then assumed, after a careful inspection of the 
data, that temperatures above the degree of the maximum rate of 
development not only did not accelerate but retarded the rate of 
development in the same proportion as an equal fall in temperatures 
below this point would retard it. 

On this supposition 84, 85, 86, 87, 88 and 89 degrees were in turn 
taken as the degree of the maximum rate of development, and cor- 
rections made in the location of the points in each case. It was 
found that 87 degrees gave the best results. Points represented by 
the crosses were then plotted, using the reciprocals of the periods 
and the average daily temperatures above 52 degrees, diminished by 
twice the average daily temperatures above 87 degrees. This 
brought all the points nearly in line with the two original points 
used in determining the line, indicating that the second supposition 
relative to the effects of temperatures above the degree of the 
maximum rate of development is correct and that 87 degrees is ap- 
proximately the degree of the maximum rate of development. 

The corrections for temperatures above 87 degrees are entered in 
column 8 and the corrected average effective temperatures (day- 
degrees) are entered in column 9. 

The day-degree is used as the thermal unit and is equivalent to 
a temperature of one degree lasting for one day. The product of 
the day-degrees above 52 degrees (column 7) by the periods (column 
4) are entered in column 10. It will be noted that they are nearly 
constant for the lower temperature, but increase as the higher tem- 
peratures are reached. 

The product of the day-degrees above 52 degrees diminished by 
twice the day-degrees above 87 degrees (column 9) are recorded in 
column 11. It will be seen that they are nearly constant for all 
temperatures. 

The average of these products is 240. 

The formula is C = P (T — 2t') in which C = constant. P = pe- 
riod in daj's, T = average day-degrees above 52 degrees and t = 
average day-degrees above 87 degrees. This formula is the equation 
of an hyperbola. Plotting the periods expressed in days against the 
effective day-degrees (T — 2t'), we have the points as represented 



322 The University Science Bulletin. 

by the concentric circles. The curve through these points is an 
hyperbola whose equation is P(T — 2t') = C = 240. 

This constant (240) is the average of 3,817 observations. We 
may, therefore, conclude that it requires an average accumulation 
of 240 effective day-degrees to bring the codling moth through the 
pupa stage. The observed variations from this average in the case 
of individuals were very great, due in some cases, no doubt, to 
errors in observation and in part to individual differences, but for 
the most part to the fact that the day was used as the unit of 
time. By using the day as the unit of time the actual period might 
in some cases be nearly a day shorter than the observed period, and 
in other cases nearly a day longer. 

In midsummer the daily accumulation of effective day-degrees 
was sometimes as high as 27, so that the accumulation of day- 
degrees for some of the observed periods might be 27 day-degrees 
greater or less than for the actual period, or the average 240. 

The following table shows the range of variations: 

Number of individuals. Day-degrees. 

1 166 

1 : 185-194 

4 195-204 

3 205-214 

110 215-224 

602 225-234 

2238 235-244 

653 244-255 

170 254-265 

18 264-275 

7 274-285 

7 284-295 

2 294-305 

Recorded accumulations of less than 205 or more than 275 were 
probably due to errors in observations; if we add 27 to the former 
and subtract 27 from the latter, we still have a variation of from 
232 to 248 due to individual differences, humidity or other causes. 
The equation for the pupal period may, therefore, be written as 
follows : 

p (T — 2t')=240± 8. 

This paper was prepared merely to illustrate the method followed 
in determining the time-temperature factors, or the equation for 
the period. The method is applicable to the investigation of any 
stage of an insect. 

If the threshold of development and the degree of the maximum 
rate of development should be found to be the same for each of 



Gli:.\.\; Ln sect Development. 323 

the stages of a given inseet, the constant of the equation for the 
wliok^ period fi-oni the deposition of the egg to tlie emergence of 
the adult may be found by adding the constants of the equations 
of the stages, and, by making suitable allowance for the time which 
elapses between the emergence of the adult and the deposition of 
eggs, a formula for the whole life cycle of the insect may be de- 
termined. 

These equations may be of i)ractical value in several ways. 

By computing the normal daily effective day-degrees in any lo- 
cality, the number of generations of the insect in that locality, and 
the normal dates on which the first eggs, or larvae, may be expected 
to appear in that place can be ascertained. If any part of the 
season should be abnormally cold or warm, the amount of retard- 
ation or acceleration in development can be computed by keeping 
a daily record of the effective day-degrees and comparing them 
with the normal temperatures. In this way we should be able to 
ascertain when the injurious phase of any insect pest is approach- 
ing long enough beforehand to enable us to take whatever pre- 
cautionary measures are necessary to avoid injury. 

In the case of the codling moth, the dates when the larvae of each 
generation may be expected to enter the fruit can be determined 
long enough beforehand to enable the owners to apply the spray 
at the right time. 

How well this method of forecasting insect injuries will work out 
in practice only time can tell ; however, the plan seems to be a 
feasible one. and one worth investigating. 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 11— October, 1922. 

(Whole Series, Vol. XXIV, No. 11.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

Some Biological Notes on Philippine Zoology, 

F. X. Williams. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4622 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 11. 



Some Biological Notes on Philippine Zoology. 

BY FRANCIS X. WILLIAMS. 

THESE few notes are very fragmentary and not all the data are 
derived from original investigations. They are set forth, how- 
ever, in hopes that the reader who has not visited the tropics will 
find something of interest in them. The writer has spent the total 
of about two years in the Philippines, chiefly at Los Baiios, Luzon, 
and thus has* become acquainted with some features of the natural 
history of this very rich locality. 

ANTS. 

It is quite evident that the ant is well off in the tropics. An 
abundance and variety of food, as well as great diversity of condi- 
tions, suit the needs of the multitude of its species and make this 
family of insects a most conspicuous one. Many ants are harmless, 
or even beneficial, while others are a great pest in the house or in 
the field. 

The well-known, red tree ant, (Ecophylla smaragdina Fab. (Cam- 
ponotinse) is occasionally a decided nuisance along the outskirts of 
the forest and in portions of cultivated districts near by. This ag- 
gressive insect has at least one thing in its favor, for while it bites 
viciously, it cannot sting. It is a tolerably large, long-legged crea- 
ture that lives on trees and bushes, where it constructs an ample 
ball-like nest of leaves spun together with the silk of its larvae, 
which are carried about by the workers as this silk is needed. 
(Ecophylla frequently swarms on tree trunks and foliage, and so a 
passer-by may unknowingly gather some up; of these a few will 
eventually and unnoticed gain a point of vantage on his person, as 
the back of the neck, and there bestow a vicious and startling nip. 
Certain homopterous bugs that produce honeydew are attractive to. 

(329) 



330 The University Science Bulletin. 

these ants and protected by them, but other insects that are not 
thus useful, and that can put up a struggle, are subjected to a gruel- 
ling treatment before they succumb. And so it is not unusual to 
come upon some unfortunate bee or beetle stretched out, more or 
less tautly, as fire fighters hold a life net, by a circle of these ants, 
each steadily and relentlessly pulling on a leg or other portion of 
the victim's anatomy. 

The large genus Polyrhachis is best represented in the forest. 
These ants also use silk in nest building, but usually mix in with it 
small debris of various sorts. A large number of the species have a 
glossy appearance. Some make a sort of ball-like nest among the 
leaves of bushes ; others will cover over a hollow, as the cut end of 
a bamboo, with a sheet of debris. Others still, often large species, 
will build on the trunks of trees, where the nests are placed in 
crevices or among the exposed roots. In these, the tubular aperture 
to the nest, with its soft, flexible rim, was found to bear fine, more 
or less inward-projecting hairs. Other nests may be high up upon 
tree trunks, and measure as much as a foot and a half long by about 
half as wide. They consist of a bulging sheet or curtain of silk, etc., 
secured along its margin to the tree trunk. I remember such a nest 
on the underhanging side of a tree with pale bark, and which color 
it somewhat resembled. As the name implies, these insects are 
armed, more or less, and sometimes in quite a fantastic manner, with 
spines or hooks, which, as they will often stick into one's fingers, 
may make capture rather awkward. Polyrhachis is only a fair 
nipper, but the offensives and defensives are effected mainly by 
raising itself upon its legs, curving the abdomen under and forward, 
and the squirting out a fluid. 

The myrmicine ants of the genera Solenopsis and Phidologeton 
can be very annoying insects, particularly during the wet season. 
Solenopsis geminata Fab., also known as the fire ant because of its 
smarting sting, is widely distributed in the tropics of both hemi- 
spheres. It is a rather small, yellowish-brown species that often 
swarm in the lowlands. In Phidologeton we have a genus of few 
species that appear more shade-loving than Solenopsis and which 
also live in very large colonies. The great majority of specimens in 
a nest are of small size, but the queens and the largest neuters or 
soldiers are comparatively immense. The ants are great travelers, 
and their irregular columns, often encountered crossing a path, are 
sparsely though conspicuously sprinkled with the great polished 
soldiers. These big, lumbering creatures are not to be feared; it is 



Williams: Philippine Zoology. 331 

the army of little workers that must be respected, for they are of a 
vindictive nature and lose no opportunity of using their sharp 
mandibles. Phidologeton will occasionally invade houses here, and 
once in a while some of the occupants are driven out of bed. There 
is a record of a village in India having to shift because of these 
troublesome ants ("Fauna, British India, Hymenoptera," II, p. 161, 
1903; Bingham quotes Rothney). The genus belongs to the Indo- 
Malayan region, and, like Sole?idpsis geminata, is largely granivor- 
ous. It is to be noted, then, that meat eaters are not always the 
fiercest. Pogonomyrmex, the aggressive and efficient stinger among 
ants in some of the more arid portions of the United States, is also a 
granivorous insect. 

While the ponerine ants are comparatively few and inconspicuous 
in the United States, this primitive subfamily is very well repre- 
sented here and contains some of the largest and commonest ants. 
Notwithstanding their superior size, in many cases they cannot cope 
against quite small, well-armed ants. This was noted in the genus 
Diacamma, and upon an invasion by such small ants, the former 
will grab bag and baggage, and, hurrying out of their nest, com- 
monly situated in a tree trunk, await the departure of the marauders. 
Besides, as is generally the case with the Ponerinse, their colonies 
are small, and sting and jaws do not count for much against a 
superior number of small ants, especially when many of the latter 
discharge a very disconcerting fluid in their battles. 

The Dorylinse include the famous driver or legionary ants of the 
American and African tropics, that in their foraging marches clear 
the path of insects and other creatures. In the Philippines I have 
observed no such formidable ants, and the only representatives of 
this subfamily familiar to me are rather small, wiry black species 
of the genus jEnictus, that travel in narrow columns. The workers, 
although blind, march with order and great activity. They appear 
to prey on other ants. 

While still on the subject of ants, it would be well to consider 
briefly some habits of the rather large muscid flies, Bengalia sp. 
These are somber-colored insects of alert habits that hang around 
the passing columns of certain ants, very often those of Phidologe- 
ton. It is quite usual to see one or more of these flies perched right 
near the moving ants, and once in a while to approach a burdened 
ant, seize and snatch away its load and to consume it at a safe dis- 
tance. Bengalia, then, feeds upon the early stages of ants and 
whatever palatable food the ants may carry. Thus, a Phidologeton 



332 The University Science Bulletin. 

army returning, laden mostly with soft, white ants or termites, was 
patronized by one of these flies. The burdens being large, however, 
necessitated several ant-carriers apiece, and thus made it rather 
difficult for Bengalia to operate. 

Among the numerous arthropods that resemble ants are various 
spiders. That some of these are more or less associated with the 
ants they resemble is beyond doubt, but the whole subject, I be- 
lieve, is still in a rather speculative stage. There are spiders that re- 
semble Q^cophylla; many, Polyrhachis ; some, Diacamma and others. 
This resemblance is often excellent, though one will learn quickly 
to differentiate ant and spider. The latter usually has much the 
better vision of the two, and so if disturbed will wheel about sharply, 
very unlike an ant. Spiders have four pairs of legs, and these ant- 
resembling species, in what suggests to us an endeavor to mimic 
antennse, will wave the first pair of legs, also unlike an ant. While 
these spiders are slender, and, like ants, properly constricted, the 
large chelae do not much resemble the jaws of ants. They have 
somewhat the habits of attid spiders. 

SOME BUTTERFLIES. 

Butterflies are not all children of the light. The tropics possess 
a number of species that are addicted to a night life, or that at least 
avoid the sunlight — mainly somber-colored insects that belong to 
the families Hesperiidae (or skippers) and Satyridse — and elsewhere 
they may be seen at sunset, or perhaps earlier on dark days, and 
sometimes also before sunrise, flying about, feeding or laying their 
eggs. I have found them coming to light but rarely, and it seems 
probable that they are not so active when the night is far advanced. 

The largest of these crepuscular species that I have observed is 
the coconut nymphalid, Amathusia phidippus Johanssen, a graceful 
brownish insect with a wing expanse of about four inches. The 
larva eats the leaves of the banana, the cocoanut, and probably of 
other palms. 

The banana leaf roller, Erionota thrax Linne, is quite a large 
skipper butterfly, whose larva makes a retreat of a strip of banana 
leaf, which it cuts away from the edge and more or less parallel to 
the midrib and rolls up as a wide ribbon. When it has outgrown or 
consumed most of this roll, it constructs another and larger one 
and pupates in the last one made. The larva is covered with a 
mealy white substance. 

A skipper even larger than the banana leaf roller, and also prob- 



Williams: Philippine Zoology. 333 

ably crepuscular, lays its eggs on the rattan (Calamus), a species of 
thorny and climbing palm. The larva is covered with a flocculent 
white material. It pupates in a neat retreat of rolled-up leaflets. 
The sensitive pupti even when gently disturbed, will so move as to 
produce what is at first a rather startling, whirring or scraping noise. 
Other large skippers feed also on palms, on plants of the ginger 
family, and on Araliaceae. 

The largest as ^\'ell as one of the handsomest of the day butter- 
flies is the bird-wing butterfly, Ornithoptera nephereus G. R. Gray, 
witli colors chiefly velvety black and brassy yellow, and a wing ex- 
panse of about six or six and a half inches. Though a fine insect, 
it is by no means the largest nor the handsomest of the genus. It is 
found in the lowlands to some distance up in the forest. The larva 
feeds on a species of Aristolochia or "Dutchman's pipe vine," which 
it shares with Papilio antiphus Fabricius, a much smaller, mostly 
black butterfly. The larvae of these two species, as well as that of 
Papilio philenor Linn of North America, resemble one another some- 
what in that they have fleshy processes on the body. The pupse also 
are swollen at the sides much more than any other pupse of Papilio 
that I know of. 

The genus Ornithoptera is sometimes considered a subgenus of 
Papilio, and ranges south, well into Australia. 

FIREFLIES. 

We have fireflies with us practically throughout the year. This 
is in striking contrast to the comparatively brief season of the adult 
beetles in the United States. But there is this much to be said in 
favor of the latter. I have seen no Luzon island lampyrid that 
equals in brilliancy the light that is emitted by Photuris pennsyl- 
vanica of the Eastern United States. 

There are quite a number of species found in this portion of the 
Pliilippines. Some have a weak luminosity, while others are quite 
brilliant, and it is a common thing to see a whole bush or crown of 
a tree sparkling like an old-fashioned Christmas tree, with hundreds 
of these insects. Such trees, especially when isolated, are visible 
from quite a distance. 

While the larvse of fireflies, being luminous themselves, are not 
difficult to find, I do not believe that many Philippine species have 
yet been associated with the adults. I have found some, as related 
by Fabre in Europe, devouring snails within their shells, the victim 
having been overcome on some bush or on the ground. In the 



334 The University Science Bulletin. 

United States the larva of the genus Phengodes is known to feed 
upon myriapods. Here at Los Bafios is a related insect with a sim- 
ilar prey. I have kept some of the larvae in captivity and fed them 
living myriapods of at least two species, and have several times seen 
the beetle larva overcome its large prey. It would grasp the myria- 
pod by an antenna, and while its victim might struggle violently, 
the sharp mandibles of the aggressor seemed eventually to pierce the 
antenna and to discharge through it a quieting fluid. At any rate, 
the prey is rendered helpless without further biting, and its interior 
hollowed out by the voracious larva. 

OTHER THAN INSECTS. 

One concerned chiefly with the study of insects cannot, however, 
fail to secure an interest in other invertebrates, as well as verte- 
brates, that come to his notice. 

The land leech is one invertebrate with which those who explore 
the mountain woods in damp weather soon become acquainted. If 
the leech feeds solely on blood, it must possess great powers of fast- 
ing. It is a tough, wormlike creature, often adorned with brilliant 
stripes, and that measures in the neighborhood of an inch in length. 
It is furnished with an anterior and posterior sucker, by the use of 
which it travels much after the fashion of certain moth caterpillars 
known as inch worms or loopers. It is very alert to a footfall, and 
from some little distance, conscious of the presence of man or beast. 
Perched on an herb or in the middle of a trail, and raised up on its 
hind sucker, it is quick to grasp a passing leg with the anterior 
sucker and climb on. Other leeches in the vicinity may w'ave their 
head end, or move along in haste in search of this prospective meal. 
Once aboard, the hungry leech seeks to make a puncture in the 
skin. This may be a difficult matter in the case of a human being 
whose legs are well wrapped in cloth puttees, or else very easy if 
leather leggings or none are concerned. Firmly affixed, it becomes 
so filled with blood as to present a very rotund and inactive appear- 
ance, and eventually to drop off. The leech, however, is on the 
whole much more disgusting than painful, as in fact one may not 
become aware of its work until it has dropped off, full fed. Like 
the hobo, the leech does not relish soap, and the barefooted native 
keeps this in mind and secures partial protection by rubbing it on. 

Most people look upon the crab as a denizen of the sea, or at least 
as an inhabitant near the seashore. However, rather small-sized 
crabs inhabit fresh-water streams near Los Banos, and I have found 



Williams: Philippine Zoology. 335 

them from two to three miles up such watercourses, the latter flow- 
ing into a large lake. One of these crustaceans was dug out of a 
decayed log along a stream at an altitude of perhaps 700 feet on 
Mount Maquiling. 

Tailless amphibians are numerous on Mount Maquiling, and may- 
occur at a considerable distance from permanent water. A quite 
large species with much of the general appearance of our tree toads 
patronizes suitable vegetation, and is not averse, on occasions, to 
visiting houses. Along the streams are large frogs which probably 
do not equal our famous bullfrog {Rana catesbiana) in average 
size. They are very neat and fearless divers, and take surprising 
headers, from some well-chosen point, into the pool, many feet 
below. 

Of lizards, skinks are the most noticeable, as they scurry away at 
your approach and rustle among the leaves. They are fond of sun- 
ning themselves along the sides of paths. Some are over a foot long 
and rather stoutly made. Very small lizards, presumably young 
skinks, are often plentiful in the forest. A species of Draco, or fly- 
ing lizard, seems to occur chiefly in the lower portions of the forest. 
Its parachuting power is secured through the extension of its ribs 
beyond the body proper, so that a wide sailing surface can be pro- 
duced. This expanded area is somewhat gaudily colored and visible 
only when the lizard is in "flight," for when at rest the ribs are 
pressed alongside the body. Then the skinny and harmless little 
creature shows nothing of its aeronautic propensities. It is arboreal, 
and sails from one tree to another in an easy descent, making a 
graceful upsweeping landing. Despite such powers, these lizards do 
not appear to wander far, as I have seen them patronizing the same 
tree for months. 

Coming to grosser lizards, the monitor lizard {Varanus salvator 
Laur.) is noticeable for its large size and noisy haste when alarmed. 
It reaches a length of several feet and is pretty well at home in the 
water, on land or up in a tree. It no doubt consumes a variety of 
food and has quite a reputation as a chicken thief. 

While snakes are common here, they are not to be found on every 
occasion, and large ones are somewhat of a rarity. Pythons {Python 
reticulatus Schneid.) occur in this region, and several years ago two 
soldiers shot a specimen about twenty-four feet long. Such exam- 
ples are few and far between. 

Of birds, the red jungle fowl {Galium gallus Linnaeus) much like 



336 The University Science Bulletin. 

some of our smaller domesticated chickens, may be heard crowing 
in the forest far from human habitation. However, they not infre- 
quently associate with the tame fowl. 

There are some very gaudy kingfishers in the woods. While some 
favor streams, others may be found in the dense woods and must 
be largely insectivorous. 

There are no large carnivorous animals here. Mount Maquiling 
has its full share of wild pigs, deer and monkeys. Any of these may 
during the night come quite close to human habitations and inflict 
minor damages to crops. Monkeys travel in companies among the 
trees and feed upon the abundant fruits. They are not especially 
noisy and their tails do not appear at all prehensile. A powerful 
but rare eagle, Pithecophaga jefjeryi Grant, is known to prey upon 
monkeys in the Philippines. 



«4 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 12— October, 1922. 

(Whole Series, Vol. XXIV, No. 12.) 

ENTOMOLOGY NUMBER V 



CONTENTS: 

Notes on Nesting of Polistes (Hymenopter.^) . . .Dwight Isely. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIYERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 12. 



Notes on Nesting of Polistes (Hymenoptera^ Vespidae).' 

BY DWIGHT ISELY, 
Department of Entomology, University of Arkansas. - 

BY FAR the most abundant paper wasp in northwest Arkansas 
is Polistes metricus Say.^ Its nests can be found almost any- 
where attached to trees or shrubbery or under the eaves of buildings. 
They are often quite numerous. As, for instance, along a stream for 
a distance of about three hundred yards the writer counted twenty- 
two nests early in May, 1921. In a small block of apple trees, 
somewhat less than an acre in extent, seven nests of this species were 
noted in October of the same year. About premises of the insectary 
of the United States Bureau of Entomology at Benton ville, Ark., the 
same season, five nests were built. 

The following notes relate largely to one of these nests which was 
built just outside of the screen, and under a shutter, of the insectary, 
conveniently situated for observation from within, where complete 
immunity from stings could be enjoyed. The history of the nest is 
as follows: 

The stem of the nest was begun May 18 by a female wasp. On 
May 20 three cells were started, and three more were begun the next 
day. As soon as the base of each, of these cells was built, an egg was 
deposited in it; thus six eggs were deposited in two days. From now 
on cells were added less frequently, but considerable time was given 
to enlarging those already started. On May 23 two more cells were 
started, and the walls of one of the first cells were 8 mm. long. Two 
more cells were added May 28. An egg was deposited in one of 
these, but not in the other until May 30. This marks the end of the 

1. Published with permission of the Secretary of Agriculture. 

2. Formerly with U. S. Bureau of Entomology: Deciduous Fruit Insect Investigation. 

3. Determined bj- Mr. S. A. Rohwer, U. S. Bureau of Entomology-. 

(341) 



342 The University Science Bulletin. 

first stage in the history of this nest. No more cells were added un- 
til near the time of maturity of the offspring. 

Hatching began May 30 with the three eggs which had been de- 
posited on May 20. Two eggs deposited on May 21 hatched on May 
31 and one on June 1. The incubation period of five eggs was ten 
days and of one eleven days. Records of incubation were not com- 
plete for the other eggs, but they required about the same time. 

The wasp larvae grew rapidly, but at an unequal rate. This was 
apparently due to the fact that some were favored in feeding. The 
first grub spun the cap, closing its cell for pupation June 9, and the 
other cells of the first-hatched larvae were closed June 1, 13, 15, 17 
and 21, and July 9, making the larval period vary from ten to 
thirty-eight days. In contrast to this extreme variation, the period 
required for pupation was exactly eighteen days in all instances, the 
wasps maturing on June 27, July 1, 3, 5, 9 and 27. All were females. 

For two days after emergence from its cell the first wasp stayed 
on the nest. After that it began going to the field, and soon was 
doing most of the field work; that is, bringing in paper for nest 
building and partially crushed insects for larval food. Caterpillars 
were apparently the most frequent prey. The parent wasp stayed 
at home, received supplies from the worker, and at first did most of 
the feeding of the larvae and actual adding to the nest. As the num- 
ber of offspring increased, there were always several on the nest 
ready to meet a field worker and unburden it of its supplies. The 
field workers also often engaged in feeding the grubs. 

Shortly before the emergence of her first offspring, on June 21, 
the parent wasp began three new cells and deposited eggs in them. 
Three more cells were started on June 27, and from then on building 
progressed rapidly. By July 10 there was a total of thirty-one cells 
in the nest, and all of the cells vacated by maturing wasps were 
renovated and eggs were again placed in them. At the close of the 
season a total of eighty-two cells had been built. 

This nest seemed to be fairly representative for rate of growth 
and size. Two other nests which were started about the same time 
were observed occasionally. On July 9 one had three wasps and 
fourteen cells; the other had six wasps and twenty-four cells. At 
the end of the season they had seventy-four and 102 cells, respec- 
tively. The total number of wasps in any of the colonies was not 
definitely known. 

The first male wasp appeared August 12, and after this for at 
least three weeks there was a large emergence of drones. The time 



Isely: Nesting of Polistes. 343 

of appearance of the first sexually mature females was not noted. 
The males did not go afield with the regularity of the females, but 
spent most of their time on the nest, so that they made up the 
greater proportion of wasps on the nest during the day. These 
males were apparently waiting on the nest for the emergence of 
sexually developed females. What was probably a preliminary to 
mating was observed twice. As a female wasp was emerging from 
her cell she was pounced on by a male and then by all the males on 
the nest. This mass of wasps fell in a ball to the ground. When 
disturbed by the writer, they separated. Mating was not observed. 
A similar performance was noted at another nest later in the month. 

Observations were not made regularly after this time, the writer 
having left Bentonville, but by the middle of October all males had 
left the nest. A large number of females still collected there, but all 
the brood had emerged and all activity had ceased. 

Considerable opportunity was offered to observe the workings of 
the sense of direction, or rather the apparent lack of any such sense. 
When the colony consisted of only a few individuals all of the wasps 
apparently depended on local observations for finding their way to 
their nest. Any change in the insectary shutters was confusing to 
them. The oldest of the workers, when returning from the field, was 
observed to alight on the shutter about a foot below and two feet to 
the north of the nest. She would run on a horizontal line below the 
nest and continue until she was about six inches past it, when she 
would turn at a sharp angle and go directly to the nest. This path 
was always followed unless the shutter was tampered with, when 
the wasp could scarcely find the nest at all. None of the other 
wasps followed this same path, but each apparently had its own 
system. 

The overwintering queens of Polistes are sometimes gregarious, a 
number of them starting a colony together in the spring. I have 
never observed them working together in building an absolutely 
new nest, but on several occasions I have seen a small number — 
never more than seven — of overwintering queens renovate a large 
nest left from the season before and start a colony together. This 
probably accounts for the very large nests of several hundred cells 
that frequently are found. These exceedingly' numerous colonies 
were much more pugnacious than the small colonies. In fact, colo- 
nies of but a few wasps are inclined to be shy rather than pugna- 
cious. 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 13— October, 1922. 

(Whole Series, Vol. XXIV, No. 13.) 

ENTOMOLOGY NUMBER V 



CONTENTS: 

Fi\'E New Species Belonging to Genus Harmolita (Hy- 
menoptera) W. J. Phillips and Fred W. Poos. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE, KAN. 



Entered at the post office at Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XI v.] October, 1922. [No. 13. 



Five New Species Belonging to the Genus Harmolita 

Motschulsky. 

{Isosoma Walker et Auct.) 

BY W. J. PHILLIPS AND F. W. POOS, 
L'nited States Bureau of Entomology, Charlottesville, Ya. 

THREE of the five species described in this paper (viz., swezeyi, 
panici and phalaridis) would fall within the genus Harmolita 
as restricted by Phillips and Emery.- Grahan^ has more recently re- 
defined the genus, however, and all five of the new species plainly 
fall within the limits as he has defined them. 

At the time this genus was revised by Phillips and Emery, the 
senior author had seen only a few specimens of H. cinnce (n. sp. de- 
scribed herewith), and since the species differed somewhat from the 
usual type of Harmolita, he did not consider it advisable to broaden 
the scope of the genus to include this species. Since that time sev- 
eral similar species have come under observation and they seem to 
be more closely related to Harmolita than to Eurytoma from a bio- 
logical standpoint, as well as from the standpoint of external char- 
acters and the character of the ovipositor. The writers fully agree 
with Gahan that it is more advisable to include these species in the 
genus Harmolita than to erect a new genus for their reception. 

The principal characters assigned to the genus by Gahan are as 
follows: Abdomen of female elongate, conical or subfusiform, with 
segments more or less subequal, the fourth segment (when prop- 
odeum is considered a thoracic segment) never greatly enlarged as 
in Eurytoma. Occiput slightly concave and immargined. Antennae 
inserted at or above middle of face; flagellum weakly clavate; 

1. Order Hymenoptera, family Eurytomidae. 

2. U. S. Nat. Mus., vol. 5.'), pp. 443-471, plates 39-48. 

3. Proc. U. S. Nat. Mus., vol 61, 1922, art. 24, p. 7. 

(349) 



350 The University Science Bulletin. 

funicle usually five- jointed and club three- jointed, though in some 
cases funicle is six-jointed and club two-jointed. Propodeum not or 
scarcely longei| than . scutellum, not sharply declivous, usually 
rugosely sculptured; with a more or less distinct median, longi- 
tudinal depression. Sculpturing of head and dorsum of thorax either 
reticulate and shining, rugulose punctate without umbilicate punc- 
tures or rugulose with a few more or less indefinite umbilicate punc- 
tures. 

Prof. C. R. Crosby, of Cornell University, kindly placed his speci- 
mens and notes on H. phalaridis at the disposal of the writers. Pro- 
fessor Crosby found that this species has two generations a year, 
each generation confining itself to Phalaris arundinacea. The re- 
maining species were reared from dried grasses collected in the field. 
No further observations on their biology have thus far been made. 

Acknowledgment is made to Mr. A. B. Gahan of the United States 
National Museum for his kindly criticism of the manuscript. 

Harmolita swezeyi, n. sp. 

(PI. XXXVI, figs. 2 and 5; pi. XXXVII, fig. 7.) 

Female. Length, 2.90 mm. Prescutum reticulately lineolate and without 
broad, shallow impressions or umbilicate punctures. Pronotal spots large, 
occupying about one-half of the anterior margin of prothorax, spots dull 
yellowish. Sometimes entire thorax and propodeum brownish. Propodeum 
without a distinct margined, median, longitudinal groove, though there is 
usually a distince median longitudinal depression. Propodeum rugulose an- 
teriorly and usually granulose in the remaining portion. Spiracular carinas 
usually distinct but weak. Abdomen equal to or slightly longer than head and 
thorax combined; conically pointed; second segment equals one-fourth length 
of abdomen; remaining segments subequal, though five is longest and three 
shortest. Legs in black specimens are black except at knees and tarsi, which 
are luteous; in brownish specimens. the legs approach the color of the thorax, 
knees and tarsi lighter. 

Antennae: Funicle five-jointed; first funicle plus ring joints longer than 
pedical; all of funicle joints usually distinctly longer than broad; middle joint 
of club apparently quadrate; scape and pedical yellowish. 

Species medium to small. Median line of face below insertion of antennae 
without markings. 

Males. Unknown. 

This species will run to the couplet separating agrostidis and websteri in the 
Phillips and Emery table of species, but may be readily distinguished from 
either species by the yellowish scape and the longitudinal depression in the 
propodeum. 

Type locality. Honolulu, Hawaii. 

Type. Cat. No. 25,471, U. S. N. M. 

Described from thirteen females reared from stems of Bermuda grass 
{Cynodon dactylon), in Honolulu, Hawaii, by Mr. O. H. Swezey. 



Phillips and Poos: Harmolita. 351 

Harmolita panici, n. sp. 

(PI. XXXVI, figs. 7 and 8; pi. XXXVII, figs. 6 and 8.) 

Female. Length. 3 mm. Prcscutvim nijinlose; pronotal spots small, occupj'- 
ing one-half or less of the anterior dorsal margin of the prothorax ; spots dull. 
Propodeum with or without a distinct continuous, median, longitudinal groove, 
though there is a distinct indication of a groove anteriorly; if groove is con- 
tinuous it is faint and very shallow, and not very clearly margined ; rugulose 
within and laterad of groove. Abdomen rather slender; longer than head and 
thorax combined; second segment comprising one-fourth to one-fifth length of 
abdomen exclusive of 1 ; 3 shortest; 4, 5 and 6 approximately the same length. 
Legs black except tarsi, knees and the lower face of the front femora, which 
are luteous. 

Antennje: Funicle five-jointed; first segment plus ring joint slightly longer 
than pedicel ; first and second segment about equal in length ; 3, 4 and 5 about 
equal in length, but each shorter than either 1 or 2; all segments longer than 
broad. Antennae black ; median line of face below insertion of antennse without 
markings. 

Species small to medium. 

Male. Length, 2.51 mm. Prescutum as in female. Pronotal spots minute, 
scarcely visible from abo\e. Propodeum without a distinct median groove; 
rugulose. Petiole granulose, somewhat shorter than hind coxse. Legs colored 
as in female except that front femora are darker. 

Antennse: Scape almost same width throughout, as seen in lateral profile, 
with no distinct shoulder near distal extremity. First, second and third 
flagellar joints with two to three annulations at distal extremity; bristles nu- 
merous and short, scarcely half the length of the segments. 

The individuals of this species without the propodial groove run to couplet 
14 in the Phillips and Emery table of species, but can be easily separated from 
hordei by having the legs not red, and from trilici by being a smaller species 
and the sculpturing being much smoother. The individuals that have the 
propodial groove run to couplet 18, in which vaginicola and secalis are sepa- 
rated. Both of these species are longer and more coarsely sculptured; vagini- 
cola has yellowish antennal scapes and the propodeum of secalis is granulose, 
which will easily separate them from panici. 

Type locality. Charlottesville, Va. 

Type. Cat. No. 25,472, U. S. N. M. 

Described from three females and one male reared from stems of Panicum 
clandestinum at Charlottesville by the junior author. 

Harmolita phalaridis, n. sp. 

(PI. XXXVI, figs. 6 and 9; pi. XXXVII, figs. 1 and 2.) 

Female. Length, 3.70 mm. Prescutum reticulately lineolate with numerous 
broad, shallow impressions; very few such impressions on pronotum, but scu- 
tellum is quite thickly pitted. Pronotal spots bright and large, occupying 
about two-thirds anterior dorsal margin of prothorax. Propodeum with a dis- 
tinct, continuous, median, longitudinal groove, which is usually margined; 
usually deep throughout, though often shallow posteriorly; numerous cross 
rugae within groove, but no indication of central carina; very rugulose laterad 
of groove; spiracular carinas prominent and spiracular area usually well de- 



352 The University Science Bulletin. 

fined. Abdomen same length as head and thorax combined; segments 3 and 5 
about of equal length; 4, 6 and 7 about equal in length and each usually longer 
than either 3 or 5. Legs: All knees, tibiae and tarsi usually reddish brown, 
femora blackish. 

Antennae: Funicle five-jointed; first funicle joint plus ring joint about twice 
as long as pedicel; first joint of funicle slender and same size throughout, the 
distal tip somewhat flaring; all of funicle joints distinctly longer than broad; 
club joints also longer than broad; antennae black and \ery slender. 

Species medium in size. 

Male. Length 2.60 mm. Prescutum as in female, but there are few thor- 
acic punctures; pronotal spots large and bright. Propodeum with or without 
a groove; groove when present is often rather poorly defined; propodeum 
usually very rugose, though it may sometimes be granulose; spiracular area 
usually well defined. Petiole usually about twice as long as broad, granulose 
and extends beyond the tip of the coxae. Legs: All knees and tarsi testaceous; 
front tibiae usually reddish brown. 

Antennae : Flagellum with peticel longer than head and thorax combined ; 
hairs on first flagellar joint approximate! j' same length as those on last joint; 
last joint bears a slender tubercle at end about twice as long as broad. Scape, 
exclusive of base, a little o\'er twice as long as broad, broadest about center, 
as seen in lateral profile ; scape as seen in lateral profile nearly twice as broad 
as first flagellar joint. There are four or more annulations at each articulation 
of the flagellum. 

This species iims to dactylicola in the Phillips- and Emery table of species, 
but the females may be separated by the following characters: H. phalaridis 
has more densely pitted scutellum ; propodeum more rugulose ; groove deeper 
and same width throughout; spiracular carinse more prominent; tibiae usually 
reddish brown; first funicle joint of antennae cylindrical, very slender and, 
distal extremity somewhat flaring at tip. All segments of antennae more slen- 
der than in dactylicola. 

Type locality. Ithaca, N. Y. 

Type. Cat. No. 25,473, U. S. N. M. 

Described from manj^ males and females reared from stems of Phalaris 
arundinacea collected at Ithaca, N. Y., by Professor Crosby and the junior au- 
thor, and from specimens reared from stems of Phalaris sp. collected at Elk 
Point, S. Dak., by Mr. C. N. Ainslie of the U. S. Bureau of Entomology. 

Harmolita cinme, n. sp. 

(PI. XXXVI, figs. 1 and 3; pi. XXXVII, figs. 3 and 4.) 

Female. Length, 3.80 mm. The whole thorax somewhat rugulose and more 
or less distinctly umbilicately punctured, the pimctures shallow and usually 
not well defined; prescutum sometimes not umbilicately punctured in anterior 
third; pronotal spots small, occupying about one-third anterior dorsal margin 
of pronotum, visible from above. Proi)odeum with a deep, margined, con- 
tinuous, median, longitudinal groove of medium width; groove with numer- 
ous cross rugae and usually with a central longitudinal carina; very rugulose 
laterad of groove; sjiiracular area usually well defined, thovigh sometimes the 
spiracular carinae are weak. Abdomen equal to or slightly longer than head 
and thorax combined, and almost as pointed as the average Harmolita; seg- 



Phillips and Poos: Harmolita. 353 

ment 2 occupies between one-third and one-fourth the length of abdomen; 
segments 3, 4, 5, 6 and 7 approximately equal in length. Legs often variable 
in color; sometimes the legs are black throughout except the knees, front 
tibiae and all tarsi, which are luteous; perhaps more often the basal third to 
half of front and middle femora and basal two-thirds of hind femora blackish; 
all tibiae, knees and tarsi almost reddish brown. 

Antennae: Funicle five-jointed; club three-jointed; first funicle joint plus 
ring jointly nearly twice as long as pedicel; all segments longer than broad; the 
first two funicle joints longest, the remaining ones of approximately the same 
length. Antennae black. Median line of face below insertion of antennae 
slightl}' elevated; doreally it appears almost carinate. 

Species medium to large. 

Male. Length, 2.90 mm. Sculpturing of thorax very much the same as in 
female, except that the umbilicate pimctures are not nearly so distinct; pro- 
notal spots verj' small, often scarcely visible from above. Propodeum variable; 
there may be a deep, rather broad, margined, median longitudinal groove, 
very rugulose within and laterad of groove, or the groove may not be con- 
tinuous and shallow, and it may be granulose within and laterad of the groove. 
In the latter case the petiole is usually granulose; when the propodeum is 
very rugulose the petiole is usuall}/^ somewhat rugulose. Petiole long, slender; 
the tip of the hind coxae often extending only to about the middle of petiole. 
Legs colored as in female. 

Antennae: Longer than head and thorax combined; scape as seen in profile 
thickened somewhat at center; no distinct club; first segment of flagellum 
approximately as long as scape; the remaining segments about of equal length; 
segments excised with about three aimulations at the distal extremity of all 
except distal segment. 

Type locality. Youngstown, Ohio. 

Type. Cat. No\ 25,474, U. S. N. M. 

Described from many females and eight males reared from stems of Cinna 
arundinacece collected at Youngstown, Ohio, by Mr. W. T. Emery and at Niles, 
Ohio, by the junior author. 

Harmolita phalaricola, n. sp. 

(PI. XXXVI, figs. 4 and 10; pi. XXXVII, fig. 5.) 

Female. Length, 3.52 mm. Prescutum somewhat rugulose, and the whole 
thorax bearing numerous but rather indefinite umbilicate punctures. Pronotal 
spots dull, minute, scarcely visible from above. 

Propodeum with a distinct, continuous, deep, medium to narrow longitudinal 
median groove; groove not distinctly margined throughout; very rugulose 
within and laterad of groove; spiracular area not clearly defined by spiracular 
carinas. Abdomen short and thick, approaching ovate; slightly shorter than 
head and thorax combined; segment 2 occupying between one-third and one- 
half dorsal length of abdomen; segments vary in length as is common in 
Harmolita, due to telescoping of segments when the insects die ; 3 and 4 often 
nearly same length; 5, 6, and 7 often about same length, but shorter than 
either 3 or 4. Legs: Basal half of front and basal two-thirds of middle and 
hind femora black ; remaining portion of legs usually reddish brown. 



354 The University Science Bulletin, 

Antennae: Funicle apparently six-jointed and club two-jointed; first funicle 
plus ring joint about twice the length of the pedicel; segments 4, 5 and 6 about 
quadrate; club joints nearly quadrate also. 

Species medium to large. 

Males. Unknown. 

Type locality. Elk Point, South Dakota. 

Type. Cat. No. 25,475, U. S. N. M. 

Described from ten females reared from stems of Phalaris sp. collected at 
Elk Point, S. Dak., by C. N. Ainslie of the United States Bureau of Ento- 
mology. 



356 The University Science Bulletin. 



PLATE XXXVI. 

1. Ovipositor of H. cinnce. 

2. Ovipositor of H. swezeyi. 

3. Propodeum of H. cinnce. 

4. Propodeum of H. phalaricola. 

5. Propodeum of H. swezeyi. 

6. Propodeum of H. phalaridia. 

7. Propodeum of H. panici. 

8. Ovipositor of H. panici. 

9. Ovipositor of H. phalaridis. 
10. Ovipositor of H. phalaricola. 



Phillips and Poos: Harmolita- 



357 



PLATE XXXVI. 




:S!* 








358 The University Science Bulletin. 



PLATE XXXVII. 

1. Antenna of the male of H. phalaridis. 

2. Antenna of the female of H. phalaridis. 

3. Antenna of the male of H. cinnoe. 

4. Antenna of the female of //. cinnoe. 

5. Antenna of the female of H. phalaricola. 

6. Antenna of the male of H. panici. 

7. Antenna of the female of H. swezeyi. 

8. Antenna of the female of H. panici. 



Phillips and Poos: Harmolita. 



359 



PLATE XXXVII. 




THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 14— October, 1922. 

(Whole Series, \'ol. XXIV, No. 14.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

The Urinary System of Phlegethontius sexta 

(Lepidoptera) G. H. Vansell. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post, office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 14. 



The Urinary System of Phlegethontius sexta Johan. 

(Lepidoptera) . 

BY G. H. VANSELL. 

(Contribution from the Zoological Laboratory of the University of Kentucky.) 

PHLEGETHONTIUS SEXTA Johan., commonly known as the 
southern tobacco worm, is used in many laboratories for mor- 
phological study material. The paper of Alvah Peterson, published 
in the September number of the "Annals of the Entomological So- 
ciety of America, 1912," treats the general anatomy of Protoparce 
Carolina Linn., but the Malpighian tubules are not shown in his fig- 
ures as fully as some other parts. If the conditions shown in the 
figures accompanying this paper differ from those found by Peterson 
in his work it may throw some light on the proposed synonomy of 
the species in question. 

Malpighian tubules are usually two in number, or in multiples of 
two, and in most cases they empty directly, or through a bladder, 
into the intestine. In the larvae of Microgaster, Koulagin found that 
they open dorsally on the outside of the body on each side of the 
anus. Those of Phlegethontius sexta Johan. empty into the small 
intestine. 

. The specimens used for dissection in this work were killed in va- 
rious ways. Some were dropped into a solution of 50 per cent alco- 
hol. 48 per cent water, and 2 per cent formalin. Immediately after 
death they were removed and the body wall split to allow the free 
passage of the preservative to the inside organs. At the time of use 
these specimens were further hardened by adding a solution of picric 
acid and chloral hydrate. Others were dropped into Bouin's picro- 
formol mixture, and later run up through alcohols. Various stains 

(36.5) 



366 The University Science Bulletin. 

were used upon that material which was imbedded for sectioning. 
Borax carmine and Delafield's hemotoxylin seemed to give the best 
results. 

The Malpighian tubules of Phlegethontius sexta are six in number 
and of a light yellowish color. Four of these are located largely 
laterodorsally to the intestinal tract, the other two being ventrally 
situated. Each tubule is apparently free at the end distad from the 
bladder. There are two bladders, located one on each side between 
the ventriculus and the small intestine (fig. 2), and just posterior to 
the ventriculus they empty separately into the small intestine 
through short, smooth tubes (fig. 3). From the opposite end of 
either bladder a tube arises which runs, dorsally and branches im- 
mediately into two; one of these tubes runs ventrally and forward, 
the other dorsally, and this one divides into two more as it nears 
the top of the ventriculus. These six tubes run forward toward the 
head and turn back posteriorly at about the second abdominal seg- 
ment, as shown in figures 5 and 6. Posterior to the bladders all 
these tubes interweave with the adipose tissue until they lose their 
identity. 

On high magnification the tubes present an interesting appear- 
ance, in that each one is covered with small nodules. These nodules 
become more numerous on the tubes the greater the distance from 
the connection with the bladder, until at the distal ends the nodules 
are very closely crowded together (figs. 7 and 8). Tracheae run to 
the tubes and branch into invisible threads upon the surface of the 
nodules (fig. 9). 

Both the nodules and the tubes are hollow, and an opening from 
each nodule leads into the tube, making a continuous cavity to all 
parts (figs. 10, 11 and 12). The exact histological nature of the 
tubes is very hard to determine on account of their delicate nature. 
The figures showing these structures are not shown here, for more 
work is being done upon them and a later paper will appear. 

LITERATURE. 

Berlese, Antonio. 1909. Gli Inset ti lore organizzazione, sviluppo, abitudini 
e rapporti coll'uomo, pp. 779-788; figs. 971, 972, 975. Societa Editrice Li- 
braria, Milano. 

FoLSOM, J. W. 1906. Entomology, with Special Reference to its Biological 
and Economic Aspects, pp. 123-124. Philadelphia; P. Blakistons & Co. 

Peterson, Alvah. 1912. Anatomy of the Tomato Larva, Protoparce Carolina. 
Annals of the Entomological Society of America, pp. 245-272; pis. XIX- 
XXI. Columbus, O. 



368 The University Science Bulletin. 



PLATE XXXVIII. 

Fig. 1. Dorsal view of the region into which the Malpighian tubules empty. 
bl, bladder; dd, Dorsal duct; ve, ventriculus; si, small intestine. 

Fig. 2. Ventral view of the same region, bl, bladder; cd, common bladder 
duct; vd, ventral duct; dd, dorsal duct; n, nodule on the Malpighian tubule; 
si, small intestine. 

Figs. 3, 4. Lateral view, showing the left and right sides of the intestine 
with the bladders and ducts, ve, ventriculus; si, small intestine; bl, bladder; 
o, duct opening into the small intestine. 

Fig. 5. Dorsal view of the ventriculus, showing the arrangement of the 
Malpighian tubules. 2nd, second abdominal segment; 9th, ninth abdominal 
segment; xx, region in which the free ends of the Malpighian tubules and 
the adipose tissue interweave into a dense mass. 

Fig. 6. Ventral view of the ventriculus, showing the arrangement of the 
Malpighian tubules. Labels as in figure 5. 

Fig. 7. Malpighian tubules under the microscojie, showing the nodules. 
This piece of the tube occurred just posterior to where a dorsal tube turns 
caudad. t, tubule; n, nodule. 

Fig. S. Section from the Malpighian tubule toward the distad end, just 
anterior to the highly convoluted area. The nodules are quite numerous here 
and the tube thickened, t, tube n, nodule. 

Fig. 9. Highly magnified portion of the tubule, showing the disappearance 
of the tracl{ae on the nodules, n, nodule; t, tube; tr. trachea'. 

Fig. 10. Cross section of tubule. 

Fig. n. Cross section of tubule and nodules. 

Fig. 12. Longitudinal section of tubule and nodule. 



Vansell: Phlegethontius Sexta. 



369 



PLATE XXXVIII. 




16 



11 



12 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 15— October, 1922. 

(Whole Series, Vol. XXIV, No. 15.) 

ENTOMOLOGY NUMBER V 



CONTENTS: 

A Brief Re.sume of Investig.ations Made in 1913 on 

Trogoderm.a inclus.\ (Coleoptera) Adolph H. Beyer. 



PUBLISHED BY THE UNIVERSITY 

LAWRENCE. KAX. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIYERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 15. 



A Brief Resume of Investigations Made in 1913 on 
Trogoderma inclusa Lee. (a Dermestid) . 

By ADOLPH H. BEYER. 

THE work herewith reported was undertaken at the suggestion 
of Prof. S. J. Hunter, who had received complaints concerning 
damage to leather horse collars. The damage consisted in more or 
less numerous perforations of the leather by some insect, which 
injury interfered seriously with the marketability of the manu- 
factured goods. It became my task to determine the insect causing 
the damage, the source of infestation, and the methods of con- 
trolling the pest. The investigations were made in 1913, but are 
reported here for the first time. 

NATURE OF THE INJURY TO THE HORSE COLLARS. 

The leather of the infested collars has the appearance of being 
perforated with numerous little round holes about the size of a pin- 
head. The insects reared from infested collars proved to be der- 
mestid beetles belonging to the above-named species. The insects 
hatch in the rye straw used in filling the collars, and the larvae live 
upon the grain left in the straw. As the larvse develop to within a 
month of the adult stage, they approach the outside of the collar 
and emerge by eating holes through the leather of the collar. 

A number of firms engaged in the manufacture of collars have 
reported damage of this character. One firm in Texas writes: 
"We wish to state that five or six years ago we had trouble of this 
kind owing to the fact that we were using rye straw with lots of 
grain still in the straw, and small worms or insects would bore 
through the leather into the collar to get at the grain. We had very 
little loss, as we soon found out the trouble, and therefore quit 
using straw that had any grain in it at all." 

(373) 



374 The University Science Bulletin. 

Two or three firms securing straw from a grower near Lawrence, 
Kan., reported serious trouble. An examination of one factory re- 
vealed fifty per cent of the collars infested. Upon examining the 
floor, walls and window and door casings, the cracks were found to 
contain many adult beetles, which in all cases were dead. The 
light and heat of the sun had a marked influence upon the distribu- 
tion and development of the insects, as the collars near the south 
windows were more seriously damaged, the injury decreasing 
gradually as one neared the north side of the building. 

SOURCE OF INFESTATION. 

Since injury was reported from manufacturers using straw grown 
near Lawrence, Kan., it appeared possible that the straw might have 
been infested before it was shipped to the factories. An investiga- 
tion was made first at the ranch where the rye straw was grown. 
It was found that the sheaves were stacked up in the barns to con- 
serve the value of the straw for collar-stuffing purposes, and during 
the winter months a specially made thresher was used in removing 
the grain from the head of the sheaf without cutting the band. After 
the rye bundles were threshed they were baled, being compressed 
and tied with wire ready for shipping to the collar factory. I found 
no trace of the insect about any part of the sheaves. They were 
apparently free in every respect from the insect. I also examined 
the threshed bundles before they were baled, and noticed that there 
was considerable grain left in the straw, which later afforded food 
for the development of the insects in the collar. I next examined 
the threshed rye in the bin, thinking perhaps the insects might be 
hibernating at this season of the year. I collected several ciuarts of 
the rye in a screw-sealed glass jar, to keep out other infestations, 
and took this, with one of the rye sheaves, to the laboratory. 

I placed some of the rye straw taken from the sheaf in a large 
glass-enclosed cylinder. This was placed in the incubator at ordi- 
nary room temperature of 70° F. I recorded the temperature each 
day and watched for the development of infestations, but noted no 
development of life for several weeks. At the end of three weeks 
I found two species of grain weevil and several parasites, which I 
removed from the jar and placed in separate vials. At the end of the 
fourth week I removed from the jar five specimens of larva.^, ap- 
parently of the same species of Dermestidse that was the cause 
of the collar infestation. April 15, the adults emerged from the 
pupa^ cases. I found them to be the same species as those causing 
the damage to the collars. 



Beyer: Trogoderma inclusa. 375 

EXTENT OF INFESTATION. 

x\s stated before, fifty per cent of tlie colhirs in one factory were 
damaged. I made a careful dissection of both sides of a badly 
infested collar by removing the leather of the facing of the collar. 
I have found only llie larval stage of the life cycle in the collars 
so far examined. T made an approximate count of all the larvae 
found in this collar, and 900 larvae was the result. I also counted 
the number of holes in the same collar leather, to get the per cent 
of damage done by each insect. The actual number of holes in this 
collar made by. this pest was 424. Approximately 1,324 of the in- 
sects infested this collar. 

The larvae varied from 2i/^ to 5 mm. in length. They were 
usually found in the heads of the rye, or eating on the scattered 
grains. The larvae eat straight across the ends or sides of the grain, 
something as a mouse gnaws, and not after the fashion of the grain 
borers. Wherever the tunneling or eating process was found, I also 
found evidence of molting. I found small larvae emerged in heads 
of rye, which is e\'idtnce that the eggs had been laid in the heads. 

LIFE HISTORY OF THE BEETLE. 

In making dissections of the collars which had been incubated 
at a constant temperature of 70 degrees for several weeks, I found 
various sizes of larvae and a number in the pupal state. None 
was found in the adult state. No eggs were found in the collar dis- 
sections. 

MATING AND OVIPOSITION. 

The beetles I found in all cases mated a day or two after emerging 
from the pupal skins. The eggs varied in number from ten to fifty. 
They were placed around indiscriminately on the bottoms of Petri 
dishes from four to six days after copulation. In a number of cases 
the female oviposited on the rye placed in the dishes, and the eggs 
adhered to the rye by means of little filaments projecting from one 
end of the egg. The young larvae hatched from eight to twelve 
days later at ordinary room temperature. The young larvae, soon 
after hatching, began to feed on the material at hand. They did 
not wander unless food was scarce or poor. I reared them entirely 
upon rye grain. The growth of the larvae depends to a considerable 
extent upon temperature and the abundance of food, and it is re- 
tarded by weather and scarcity of nourishment. The foregoing 
factors are not, however, the only causes of slow development. 
I noted in my experiments that the growth of specimens varies 



376 The University Science Bulletin. 

under identical external conditions. Very often a number of the 
specimens attain full size, metamorphose, and produce young long 
before others of the same generation. The majority, however, ma- 
ture in about five months. 

THE EGG. 

The eggs are about % mm. in length and about Yq mm. in width. 
They are oblong in shape and slightly arched. Not uncommonly 
they were found adhered together in pairs. One end of the egg 
usually has hair or threadlike projections. Each egg has a number 
of ridges running lengthwise and crosswise (see plate XL). The 
eggs are whitish in color, translucent, and the surface appears rough, 
and is of delicate skin covering the aqueous interior. It is easily 
broken. The filaments described at the end of the egg adhere to 
any object with which they come in contact. 

MOLTING. 

I found without exception that there is much variation in the 
rate of molting and the number of larval skins shed by the different 
individuals of this species. Under normal conditions the larvae molt 
twice in about every two weeks. Many peculiarities are worthy of 
mention. The same specimen often sheds its skin irregularly, some- 
times within ten days, and again, under the same conditions, not 
until a period of three or four weeks has lapsed. The rapidly grow- 
ing individuals molt more frequently than do those which have 
about attained their full size. The specimens that are slow in de- 
velopment molt less frequently than do the larvae which develop 
at the average rate. The full-grown larvae previously spoken of, 
which continue to live for a long time before entering the pupal 
stage, have a decidedly slow rate of molting. The average rate is 
about once in every three weeks, and there is a gradual decrease 
as the specimen grows older. 

Thus I have found in all my observations that the number of 
molts is by no means constant. As previously mentioned, the ma- 
jority of specimens which complete their life history in about five 
months, shed their skins from ten to fifteen times, whereas many 
of the individuals with the prolonged larval history molt as often 
as twenty times. 

The larva? never eat their own skins, nor the skins of other indi- 
viduals of this species, even though they may be in the most extreme 
stage of starvation. This was conclusively proven by placing speci- 
mens in a glass vial for the purpose of starving the larvae, and after 



Beyer: Trogoderma inclusa. 377 

several months of starvation, during which the larvae had molted 
several times, the skins were never attacked. This fact was also 
proven in the collar dissections which I made. In one collar in 
particular, which had been kept in stock for three years after the 
infestation was noticed, nearly all' of the rye seed had been de- 
voured, and the straw and inner surface of the collar were literally 
full of molted skins, and a large percentage of the larvae had escaped 
from the collar. 

Just before molting the specimens become inactive, and a break 
appears in the larval skin along the median dorsal line. This ex- 
tends from the head along the thorax and partly down the abdo- 
men (see plate XL). The larva assumes a semicircular position, 
which permits the extrication of the thorax and the head. The legs 
are then pulled out of their covering, and the light-colored larva 
crawls out of the exuvia. Its soft covering soon hardens and be- 
comes chitinous, and within a few hours assumes the natural yel- 
lowish-brown color. 

PUPATION. 

When the larva reaches full growth the pupa begins to form 
within the last larval skin. This is noticeable by the size, shape and 
the lack of movement or locomotion. The pupa is slightly shorter, 
larger in diameter, and apparently makes no movements. Four or 
five days later the skin splits down the median dorsal line and the 
light-yellowish pupa is exposed. The period of molting lasts from 
ten to twenty days at ordinary room temperature. When the insects 
are fully developed they emerge through the large dorsal opening 
of the pupal skin. Should a specimen be forced out of the larval 
case when not fully matured, though capable of locomotion, it in- 
variably returns to its former position within the protective larval 
skin upon coming in contact with it. The pupae upon emerging 
are of a whitish color ; then comes the darkening of the hair on the 
thorax and elytra. In three or four days the thorax and elytra take 
on a reddish color, commencing at the thorax and shading back. 
The female remains in the pupa case a day or two longer than the 
male. The average life of the adult is about eighteen days. 

FOOD HABITS. 

I have found that this species can subsist upon a large variety of 
substances. In considering the relative value of some of the sub- 
stances as food for the larva, I found that the pest apparently 
thrives best on cereals. As was formerly stated, the larvae were, in 
their natural state, in almost every instance found living on the 



378 The University Science Bulletin. 

grain in the horse collars. Where the grain was most abundant 1 
found the largest percentage of larvae. In wandering in search of 
food many holes were made through the straw, upon which they fed 
in the meantime ; and upon coming in contact with the inner surface 
of the leather, holes about the size of a pinhead were made, through 
which they emerged, thus injuring the salability and market value 
of the leather goods. I placed a number of the specimens, soon 
after they were hatched, on a leather diet, and I found they did not 
eat at all. 

Mr. J. E. Wodsedalek (1912) says with reference to the species 
Trogoderma tarsale: 

"A number of the specimens were placed on a feather diet, and although 
they are now two years old, they have grown but veiy little. When they were 
one year old they were veiy little larger than the newly hatched individuals, 
and at the end of the second year of life they reached a meager size equal to 
that which specimens fed on insects ordinarily attain in two weeks. Their de- 
velopment on wool is even slower." 

F. H. Chittenden (1897) states: 

"One jar of flaxseed from the museum department is infested chiefly by this 
common museum pest. Many of the larvae may be seen through the glass, and 
large patches of their yellowish-brown gnawings and excrement show whene 
they have been at work. In castor beans a few were present. 

"That this species of Trogoderma can subsist on a vegetable diet is as posi- 
tive as it is surprising. No other Coleoptera, to my knowledge, live on oil 
seeds, and I had nearly arrived at the conclusion that this form of matter was 
the nearest approach to animal food available, and that these insects could 
only thrive on such vegetable substances as contain a considerable portion of 
oleaginous matter. Judge my astonishment when a few weeks after the dis- 
covery of the Trogoderma living in oil seeds, Doctor Howard brought me a 
box nearly full of cayenne pepper in which were several Trogoderma larvae. 
The most careful search failed to show even a fragment of that well-known 
red pepper pest, Sitodrepa panicea, or of any other insect than the Derme- 
stida?. Subsequently the adult was reared and proved to be Trogoderma tar- 
sale. 

"To ascertain whether this species would breed on so pungent a substance 
as cayenne pepper, a few adults were confined with a quantity of this condi- 
ment. In due time larvae appeared, and when examined, August 20, or nearly 
ten weeks from the time the eggs were deposited, were in vigorous condition, 
the average individual measuring a tenth of an inch in length, or about half 
that of the full-grown larvae. Toward the end of September, while passing 
through the museum of this department, my attention was attracted by an 
accumulation of powder and dust about the edges of an exhibit of peanut oil 
cake, and another of Indian-turnip bulbs. A large number of the larvae and 
their cast skins were found under the cakes, also in the flour and meal prepared 
from peanuts. The Indian-turnip bulbs were very old and dry, and might have 
been on exhibition twenty years or more. 



Beyer: Trogoderma inclusa. 379 

"When this irijioct infests a siibstance of similar color and consistency to 
Hour and meal, only a few laiv;^. are sufficient, on account of their extraor- 
(.linary habit of frequently molting, to occasion alarm. In fact, appearances 
are much worse than the reality. Thus in a small jar of peanut meal in which 
these larv* had taken up their abode, about forty larval skins had accumu- 
lated when examined Sejitembcr 27. completely covering one-half of the sur- 
face of the meal, and giving the impression of a whole colony of insects. 

"While the division of entomology was moving into new quartere a bag of 
Saskatchewan spring wheat, formerly kept in stock for distribution, and de- 
scribed on the label as a hard, amber variety with an exceedingly heavy grain, 
was unearthed, in which the lan'se of this insect was livdng, three being present 
and no other insects except a colon\- of Arithrenus and a single stray Silvanus. 
In fact, this grain is so hard and flinty that weevils would not flourish on it. 
Soon afterwards I found larva; in another lot of wheat infested with Silvanus 
and in corn containing Calandra crryza and other small beetles. About the 
same time, Mr. Frank Benton brought me some larvae found in beehives, 
where they apparently fed upon propolis in bee glue. There are sexeral re- 
corded instances of Dermesies lardarius feeding upon wax, or, more properly 
speaking, honeycombs, and it is therefore fairly certain that Trogoderma has 
the same habit, although not previously reported in beeliives. 

"Among the divi.sional notes I find one recording the receipt of six larvae 
of this species in a box of red pepper from a correspondent in Utah, Novem- 
ber 22, 1882. These lar\a? were kept in a box of red pepper for a year, at 
which time fifty-four cast skins were noticed. The box was examined January 
14, 1887, or over four yeare from the time of its receipt, when two larvae and 
seventy more cast skins were found, but no trace of beetles, although it had 
been kept closed so that it was impossible for either laiTse or adidts to escape. 
It is very ob^■ious that four larvae, or the beetles that develoi)ed from them, 
had died in the interim and were devoured by their fellows. In any case, the 
achdt was not reared, and no published statement was made of the lar\-8e hav- 
ing been foimd living in the condiment. 

"The capability of this species breeding in other seeds was demonstrated 
by the discovery of the larvae living upon 'kolu,' an edible leguminous seed 
somewhat resembling a cowpea. The insect had evidently been first attracted 
by the dead bodies of the original inhabitants of the seeds, the weevil, Bruchus 
chinensis, but had afterward fed upon the seeds, even hollowing them out and 
leaving only the empty shells. In a similar manner, larvae were found, to- 
gether with those of AttayenuH, in millet and pumpkin seeds that had formerly 
been inhabited by the polyphagous Indian meal moth. Plodia interpunctella. 

"In the case of the six lan-ae found in the red pepper, it is not likely that 
four of them metamorphosed, because if they had it is certain they would 
have been devoured by their fellows. The hard, chitinous covering and the 
elytra are never completely devoured, even by star\'ing specimens. It is much 
more pi-obable that they died in the larval stage, and were later de\oured by 
the other two larvae, or they might have shriveled up and darkened, and were 
thus easily overlooked. That the two larvae which were present four years 
later were two of the original six is highly probable. There are several larvae 
in our laboratory which were obtained three years ago. when they were full 
grown, and they have not changed any since." 



380 The University Science Bulletin. 

behavior. . 

Naturally the larvae manifest a strong negative reaction to light, 
and make effort immediately after hatching and when disturbed to 
seek a shaded or other place of concealment. If placed near a light 
or window they soon begin to crawl away from the light. It is 
also quite noticable when the specimens are placed in a dark room 
and a strong light is introduced at one end of the glass dish con- 
tainer. This negative phototactic reaction persists throughout life. 
It is at its highest sensibility to light just before pupation. Thus 
pupse are most frequently found in shaded or dark places which 
afford them a favorable means of protection. The adults, both male 
and female, retain their negative reaction to light after emerging 
from their pupal skins. During the period of sexual excitment which 
follows a day or two later, the insects are still negative, and the 
females remain decidedly so until their eggs are safely deposited. 
Several hours or a day after egg-laying, they gradually become 
indifferent to light, and finally a complete reversal of their former 
reaction follows. The males also become positively phototactic 
during the last days of their lives. The larvae of all stages feign 
death upon being disturbed. However, when disturbance is con- 
tinued from a few seconds to a minute at the most, they no longer 
respond in the same manner. When disturbed the adult insects 
make themselves very compact by drawing thorax up close to the 
rest of body. The head is drawn upward and under the thorax, legs 
and antennae are folded up, and death is feigned a considerably 
longer time than in the larval state. The average feint lasts from 
one to ten minutes 

VARIATION IN SIZE. 

The adult male insects are smaller, as a general rule, than the 
female insects, but the small individuals are not always males. 
There is much variation in the size of the adults. They are from 
1.5 mm. to 4 mm. in length, the width also being proportionate. It 
is difficult to determine the exact cause for this variation in size. 
Poor nutrition evidently has effect upon the size. However, small 
individuals appear among the large ones which have lived under 
very favorable conditions. I noted marked variations in the size of 
the different larvae of the same brood within a day or two after 
hatching. I observed the fact, however, that the small, slowly de- 
veloping larvae do not always produce small adults. 



Beyer: Trogoderma inclusa. 381 

adaptation to food supply. 

An interesting phase of the study of the life history was the ex- 
tremely long period of time the larvae can sustain themselves with- 
out food. I placed forty larvae in Petri dishes^ ten representative 
stages, varying from newly hatched to full-grown individuals, with- 
out any food whatsoever, for the purpose of determining the period 
required to produce starvation. I also added a number of Petri 
dishes, each containing one larva ranging from 1 to 6 mm. in length, 
and another Petri dish was added, containing a number of a definite 
size, to determine whether they would eat their skins or not. Meas- 
urements were made of all the individuals and records kept. Dishes 
were examined regularly and measurements made of the representa- 
tive stages. I also made a record of the cast skins. I found that 
the larvae never devour the molted skins of themselves or other speci- 
mens. I detected no evidence of cannibalism among the larvaB, even 
the full-grown starving specimens never attacking the much smaller 
individuals. Practically all of the larvae shed their skins shortly 
after being deprived of food, but the molting process from this on 
was very much slower. The measurements showed in all cases that 
the different larvae decreased in size about one-half their normal 
length after eight months of life at a temperature of 70 degrees and 
in ordinary daylight. The newly hatched began to die when about 
three months old. The larvae of the middle stage up to the adult 
stage were all still surviving, and judging from existing circum- 
stances, the survival, especially of the full-grown larvae, would be 
considerably over one year. Experiments were carried on with 
reference to different kinds of diets or foods taken from the collar, 
as rye seed, rye straw and leather, and in drawing comparisons it was 
found that they thrived and grew rapidly upon the rye seed, but 
refused any of the other materials contained in the make-up of the 
collar. 

CONTROL MEASURES. 
HEAT. 

I first took the trouble to look over the field of available literature 
relative to the control of this class of insects. I found that the 
French were the first to know the value of heat, and to devise con- 
trivances for the heating of infested buildings. Experiments were 
made by Professor Webster to ascertain the amount of heat required 
to destroy the Angoumois grain moth, which gave the following 
results : 



382 The University Science Bulletin. 

"A temperature of 140 degrees continued for nine hours literally cooks the 
larvse or pupse, a temperature of 130 degrees for five hours is fatal, as is also 120 
degrees for four hours, while 110 for six hours was only partially effective." 

It was also found that wheat could be subjected to a temperature 
of 150 degrees for eight hours without impairing its germinating 
properties. In the second report of the state entomologist of New 
York, Prof. J. A. Lintner, speaking of Tribolium jerrugineum infest- 
ing grain and flour, says : 

"A naoderate degree of heat, 120 to 130 degrees, continued for a few hours, 
would in all piobability suffice to kill all the eggs, larvse and pupse in the 
material, while a higher temperature, perhaps 150 degrees or more, would be 
needed for the beetles." 

Professor Chittenden, in his paper on "Insects Injurious to Stored 
Grain," states: 

'"Prior to the adoption of carbon disulphide as a fumigant, heat was relied 
upon in the destruction of these insects. A temperature of from 125 degrees 
to 140 degrees Fahrenheit continued for a few hours is fatal to grain insects, 
and wheat can be subjected to a temperature of 150 for a short time without 
destroying its germinating power." 

A large number of the experiments of this nature were made rela- 
tive to the discovery of a method to destroy grain moth, and from 
the results of these experiments many of the grain insects could 
probably be destroyed in the same manner, but it would require a 
higher temperature to destroy the adults than the larvae or pupse. 

In the first experiment about thirty of the adults and larvse were 
placed in a Petri dish which contained rye seed. A thermometer 
was placed in the vial, with the bulb resting in the middle of the 
rye, in the Petri dish containing the rye grain and various stages of 
Trogoderma inclusa. The Petri dish was then placed in a dry-heat 
oven. The bottom and surface of the interior of the oven was 
covered with asbestos, and the Petri dish was placed upon it, to 
allow uniform heating. The heat of the oven was raised to 86 de- 
grees Fahrenheit before proceeding with the experiment. No change 
was noted in the action of the insects. At a temperature of 100 de- 
grees the adults and larvae began to crawl out of the grain; at a 
temperature of 110 degrees both larvse and adults manifested ex- 
cessive excitement, and were making every effort to escape. They 
continued to be cjuite active until the temperature of 115 degrees 
was reached. At this temperature the adults and larvse became less 
active, and at a temperature of 118 degrees the adults were all dead, 
and also a large number of the larvse. At a temperature of 119 
degrees there was no sign of life. To be sure that I had killed all of 



Beykk: Trogoderma inclisa. 383 

the insects, I raised the temperature to 120 degrees, and then the 
insects were removed and pUiced in an incubator and given a chance 
to recover, but the test showed finally that they were all dead. I 
repeated the experiment a number of times, and found that as soon 
as a temperature of from 119 to 120 degrees was recorded it proved 
fatal to all stages of the insect. It required about thirty minutes 
to reach this temperature. 

In a second series of experiments I continued to use the dry-heat 
oven, again raising the temperature to 86 degrees. I then took one 
of the infested horse collars and made a hole in the leather, through 
whicli I inserted the bulb of a thermometer into the interior of the 
stuffing. The collar was then placed in the oven on an asbestos floor 
to allow equal heating. The temperature was gradually raised to 
120 degrees, which took about ten hours. After leaving the collar 
in the oven for a period of ten hours it was removed and placed in 
the incubator to allow a chance for the insects to develop again. A 
day or two following I removed the collar and proceeded to make 
a thorough dissection of it. I found that the larvae and adults were 
all dead. Upon further incubation of the straw stuffing it was found 
that the eggs had also been destroyed by the maximum temperature, 
as no more of the insects were hatched. 

After demonstrating in the laboratory by experiments that this 
species of insect could be destroyed at a temperature below that 
which would be injurious to the leather of the horse collars, steps 
were taken to test out its results as to the practibility in a seriously 
infested collar factory overrun with Trogaderma inclusa, of the 
results which were demonstrated in the laboratory. 

The means of extermination was left in the hands of the experi- 
menter. The collar warehouse w^as the only infested floor, and it 
was located on the sixth floor of the factory. The heating facilities 
were too inadequate to produce the temperature required to ex- 
terminate the infestation; hence, according to instruction, a small 
room, twelve feet long, eight wade and twelve high, was constructed, 
and lined on the interior with asbestos. Two large steam-heat radi- 
ators were installed, and the collars hung on brackets about the 
room. One large thermometer was placed in the room and several 
smaller ones inserted in the collars. The results of this work proved 
successful. 

In this experiment also two thermometers were employed. One 
was placed in a collar and the other in the chamber. By recording 
the temperatures at short intervals and plotting them on a chart, 



384 The University Science Bulletin. 

it was evident that the temperature rise in the collar lagged behind 
that of the chamber. This lag increased as the temperature rose 
until at 122° F 'f here was a difference of nearly two hours. Thus 
the steam was turned into the radiators at 8 a. m. and while the 
chamber temperature reached 122° F. at about 4 p. m. the collar 
temperature did not reach this until 6 p. m. 

COLD. 

A temperature control machine such as is used by this depart- 
ment was used to demonstrate the possibilities of a freezing tem- 
perature as a factor in the means of exterminating the pest. A 
number of the specimens were placed in a vial in the temperature 
machine. The temperature was reduced to two degrees below zero 
and held constant for ten hours. The larvae were apparently dead, 
but after a short period of incubation the specimens began to be 
active again. I did not continue the experiments, as I realized the 
impracticability of this means of extermination. 

CARBON DISULPHIDE. 

Carbon disulphide is a foul-smelling liquid that volatilizes readily 
at ordinary temperature, and produces a heavy vapor that is 
deadly to insects of all kinds when they are confined in a closed 
space and must breathe it. In reading the data of the various books 
and bulletins at hand on fumigation, it was found that this gas 
is especially useful against species infesting stored grains and mills, 
etc., overrun with Dermestidse, moth, and so on. There was reason 
to believe, then, that the insects within the collars could be ex- 
terminated if the carbon disulphide could be properly administered 
to the interior of the collar. 

I made a carbon disulphide extermination test of the Trogoderma 
inclusa as they occurred in the horse collars. I prepared a box 
for inclosing one of the collars by covering it with paper on the 
inside, and closing it with a tight cover. A small amount of carbon 
disulphide was injected into the collar by means of a specially 
devised syringe. The syringe used is modeled after the ordinary 
type with the exception that a nozzle about four inches in length 
was devised to reach all parts of the interior of the collar. The in- 
strument is made of steel for durability and strength, as consider- 
able force is required in making the insertion. The nozzle of the 
syringe was inserted in the collar stuffing, entrance being made be- 
tween the seams; thus there was no injury to the collar. It was 
injected at spaces of about four or five inches apart. From six to 



Beyer: Trogoderma inclusa. 385 

eight drams were injected into each collar. The collar was then 
inclosed in the box and left for twenty-four hours. After making 
a thorough dissection of the collar, the various larval stages found 
were all dead, and upon incubation no evidence of recovery was 
noted in any of the larva? or adults. 

HYDROCYANIC-ACID GAS. 

Hydrocyanic-acid gas is a vapor very destructive to all life. The 
gas is produced by adding potassium cyanide to sulphuric acid. I 
used these in the following proportions: 

Potassium cyanide, 98 per cent pure 1 oz. 

Sulphuric acid, specific gravity 1.83 2 oz. 

Water 4 oz. 

I placed several of the infested collars in a closed chemistry hood. 
I put an earthen vessel inside, containing water, poured the sul- 
phuric acid slowly in the water, and then added the potassium cya- 
nide, and immediately closed the hood and left the collars exposed 
to the hydrocyanic acid for two hours, and then opened the hood 
and let the gas escape. I then took the collars from under th-e 
hood and examined the stuffing. There was no trace of life or re- 
covery of the insects in the different stages. 

SUMMARY AND CONCLUSION. 

The Trogoderma inclitsa discussed in this paper were found in 
horse collars that were sent to this department, requesting our ad- 
vice and assistance in determining the kind and source of infesta- 
tion and measures to be used in the control of this damaging insect. 

With regard to the source of infestation: Upon opening the col- 
lars and finding grain in the straw stuffing, and upon examination 
of the grain in many instances, I noted that it had been eaten upon 
by the larvae of this insect, and through the instrumentality of my 
series of investigations in relation to the life history, I found that 
they thrived much better upon the grain than any other material 
found in the make-up of the collars. This gave me a clue as to the 
source of infestation. After getting some of the stored unthreshed 
rye, and some of the rye seed which had been stored where it had 
been raised, I found that a series of incubations produced exactly 
the same species that was found in the collars. 

In considering the matter of infestation and the measures to be 
used for its control, I suggest that the grain be eliminated as nearly 
as possible from the straw to be used in the stuffing of the collars, 
and there will be no liability of serious infestation. However, as I 



386 The University Science Bulletin. 

have experienced personally and through correspondence; many of 
the people engaged in the manufacture of horse collars are not aware 
of the value of clean-threshed straw, the seed of which forms an 
abundant food supply for these insects, and thus follows the dilemma 
caused by the infestation. Applicable to such cases, I have per- 
formed a series of experiments to demonstrate their relative value 
concerning the means of extermination. Experiments were carried 
on with heat and cold temperatures and carbon disulphide and 
hydrocyanic-acid gases, and in conclusion I feel at liberty to state 
that heat is to be preferred as the best means of control. It insures 
extermination, and is by far the most economical as well as the 
safest means of eradicating the infestations. 

With reference to the matter of multiplicity and distribution of 
this species of insect, it was found to be a rare species generally 
distributed over the the United States and Europe, and is omniv- 
orous in its feeding habits. I found this species to thrive and de- 
velop much more rapidly upon seeds and grain foods than any other 
available material which I had at hand for testing out the food 
habits. 

In making a summary of the variations of the life history of a 
number of individuals of the same generation, I noted: 

1. The adults oviposit from four to six days after emergence. 

2. The number of eggs laid by different individuals varied from 
ten to forty-five. 

3. The eggs hatch in from eiglit to twelve days at ordinary room 
temperature. 

4. The larval life lasts about five months, on the average. 

5. The time of pupation is from ten to fourteen days. 

6. The adult lives from eight to twenty-five days. 

BIBLIOGRAPHY. 

F. H. Chittenden. 1893. — Herbiverous Habits of Certain Dermestids. Bull. 

2, N. S. Div. Ent., U. S. Dept. Agr., i)p. 36. 37. - 
1897. — Granivorous and Other Habits of Certain Dermestids. Bull. 8 N. S. 

Div. Ent. U. S., pp. 14-24. fig. 1. 
L. O. Howard. Extract from Corres. Bull. 44, Div. Ent, U. S. Dept. Agr., Apr., 

pp. 90-99. 
H. F. Jayne. 1882. Revision of the Derm, of the U. S. Proc. Amer. Philos. 

Soc, vol. XX. 
C. V. Riley. 1883.—Trugoderma as a Museum Pest. Amer. Nat., vol. 17 

p. 199. 
1S83. — Number of Molts and Length of Lar\-al liife as Influenced by Food. 

Amer. Nat., vol. 17, pp. 347-548. 



Beyer: Trogoderma inclusa. 387 

F. H. Snow. 1SS2.—A New Museum Pest. Psyche, vol. 3, pp. 351, 352. 

1S94.— Insect Life, vol. VI, p. 226. 

1894- — Proceedings of the Columbus Hort. Soc, vol. IX, p. 12; Apr. 

.?S56.— Canadian Entomologist, vol. XXVIII, p. 262; Oct. 
W. S. Blatchley. Coleoptera of Indiana, p. 593. 

Dec, 1912, Annals of Ento. Soc. of Amer., p. 367. 
Le Conte. Synopsis of the Dermestidse of U. S. Proc. Phil. Acad. Nat. Sc, 

vol. VIII, 1854, pp. 106-113. 
Casey. Review of the American Dermestidse. Jour. N. Y. Ento. Soc, VIII, 
1900, pp. 138-165. 



388 The University Science Bulletin. 



PLATE XXXIX. 

Photographs of leather horse collars damaged by the dermestid Trogoderma 
inclusa Lee. In figures A and B, white-headed pins were inserted to indicate 
the position of the holes made by emerging beetles. Figvu'e C shows the exit 
holes made by the beetles. 



PLATE XXX IX. 





B 




(389) 



Photographs by P. A. Readio. 



PLATE XL. 

Truyodeniia inclusa I^ec. 

1. Ventral view of beetle. 

2. Dorsal view of beetle. 

3. Ventral view of jnipa. 

4. Eggs. 

5. Antennae. 

6. Ventral view of larva. 

7. Pupa. 

8. Dorsal view of larva. 

(390) 



Beyer: Trogoderma inclusa. 



391 



PLATE XL. 








.-(rfS 





'V'hi> 8 



THE 

KANSAS UNIVEESITY 

Science Bulletin 



Vol. XIV, No. 16— October, 1922. 

(Whole Series, Vol. XXIV, No. 16.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

The Larva of a Chironomid (Diptera) P. W. Claassen. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS INIYERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 16. 



The Larva of a Chironomid {Trissocladius equitans n. 
sp.) Which Is Parasitic upon a May-fly Nymph (Rith- 
rogena sp.).* 

BY p. W. CLAASSEN. 

IN AUGUST, 1919, while spending a few weeks in Colorado, the 
writer was collecting aquatic insects in the Big Thompson river 
in Estes Park. This river is a typical mountain stream. The water 
is very cold and rushes along in a swift current over a stony bed. 
Aciuatic insects of the swift-water type were very abundant. Among 
the May-fiy nymphs collected there were found about a dozen speci- 
mens of one species which presented a curious appearance. Upon 
examining some of these nymphs it was found that each one carried 
upon its back a large, white dipterous larva. These larvae had at- 
tached themselves to the thorax of the nymphs underneath the wing 
pads. The large size of the larva forced the wing pads of the May- 
fly nymph upward at a very decided angle and gave the nymph a 
humped-up appearance as it rested upon the stone. 

All the collected material was preserved in alcohol and taken back 
to Ithaca, N. Y., for study. When the material was examined more 
closely in the laboratory, some of these May-fly nymphs were found 
to carry larvae, while others of the same species carried pupae of the 
dipterous insect. 

The May-fly nymph proved to be a species of the genus Rithro- 
gena. Although it was impossible to determine definitely the genus 
of the diptcron, Dr. 0. A. Johannsen, who examined it, felt certain 
that it belonged to the family Chironomidse. The scarcity of mate- 
rial and the lack of adult forms, however, made it impossible to sat- 

* Read before the joint session of the Entomological Society of America and Ecological 
Society of America, January, 1922. Withdrawn for publication in this bulletin. 

(395) 



396 The University Science Bulletin. 

isfactorily establish the relationship which existed between these 
two forms. 

During the summer of 1921 the writer made another trip to Estes 
Park, Colo., and again found these May-fly nymphs in the same 
situation in the Big Thompson river where they had previously oc- 
curred. A careful search was then made over a distance of a mile or 
more in this stream, but there was only this one particular spot in 
which the Rithrogena nymphs could be found in considerable num- 
bers. This was in a part of the stream where the current was quite 
swift and the water not over six to ten inches deep. The bed of the 
stream was covered with stones, many of the stones projecting 
above the surface of the water. May-fly nymphs, stone-fly nymphs, 
caddis worms and other swift-water forms were very plentiful. As 
many as five or six specimens of the Rithrogena nymphs occurred on 
a single stone. Upon taking a stone out of the water and turning it 
over, these nymphs would quickly glide to the under side of the 
stone, appearing to be much more elusive than any of the other spe- 
cies of the May-fly nymphs present. A total of nearly 300 of the 
Rithrogena nymphs were collected, and more than ninety per cent 
of them were found to carry either a larva or a pupa of the chi- 
ronomid. 

An attempt was then made to rear to the adult stage the May-fly 
and the chironomid. In order to accomplish this a number of the 
nymphs which carried pupsB were placed in small wire cylinder 
cages. Some of the cages were placed in the part of the stream 
where the nymphs naturally occurred, while others were placed in 
a spring near the writer's cottage, where they could be kept under 
close observation. Three males of the chironomids emerged on 
August 7, two of them from cages in the stream and one from a cage 
in the spring. A female also emerged on the same date, but it es- 
caped. On August 8 two males of the May-fly emerged, one from 
a cage in the stream and the other from a cage in the spring. On 
the previous day two females of the May-fly were caught in a net 
near the same spot where all the material was collected, and these 
later proved to belong to this same species. 

The necessity of leaving the park on August 8 prevented any fur- 
ther rearing work, and the material from the cages was added to the 
alcoholic specimens and taken back to the laboratory at Cornell 
University. 

The May-fly has been determined by Dr. J. G. Needham, who 
recognizes it as a new species of the genus Rithrogena. A descrip- 



Claassen: Larva of a Chironomid. 397 

tion of this species will be given by Doctor Needham in a paper 
which is soon to be published. 

The chironomid is a new species of Trissocladius, a genus new to 
this country.* 

DESCRIPTION OF THE STAGES OF THE CHIRONOMID. 

Trissocladius equitans n. sp. 

ADULT. 

Length, 4.5 mm., exclusive of antennae. General color, blackish brown. 

Head blackish ; ej-es black, naked, slightly emarginate on inner margins, the 
facets moderately rounded; distance between the eyes from above greater 
llian the length of the eye. Labium short, thick, and reaching to the second 
segment of the palpi. Palpi short, three-segmented; the first segment nearly 
twice as long as broad; second segment twice as long as broad; third seg- 
ment graduallj' tapering to the tip and a little longer than second segment; 
total length of palpi about .25 mm. Antennae composed of fourteen segments, 
the terminal one-half again as long as the combined length of segments two 
to twelve; ba^al segment large; second segment about twice as long as third; 
total length of antenna L3 mm.; entire antennae covered with long hairs, the 
basal hairs as long as the terminal segment. Epistome without bristles; trans- 
verse suture distinct. 

Thorax blackish brown, with whitish pruinose patches; dorsum with a 
naiTow median line, depressed in front and raised into a feeble carina behind 
the middle; surface smooth except for a few hairs on each side of the broad, 
flattened longitudinal area of the mesonotum ; collar incised in middle, angles 
rounded; scutellum and metanotum blackish, smooth; plura and pectus black- 
ish, smooth. 

Abdomen compressed, blackish, with long, yellowish hairs ; basal segment of 
the male clasper about twice as long as the distal segment, spoon-shaped, 
the median projection or tubercle blunt, with short hairs, basal segment below 
with long hail's; distal segment narrow at base and gradually enlarging toward 
the apex, a few hairs above and underneath with two short spines near 
the tip. 

Wings milkj^ white, reaching to the base of the claspers; surface finely 
punctate; anal lobe produced and fringed with long hairs; venation as in 
figure 14. 

Halteres pale, slightly infuscated. 

Legs light brown, hairy; tibia of front leg L4 times the length of metatarsus; 
a single distinct spur at distal end of tibis. the ones on the posterior legs 
being much larger; fourth and fifth tarsal segments of equal length; tarsal 
claws squarely truncate, surface fluted; no pectinate empodium present. 

Type in the Cornell University collection, now mounted in balsam. Para- 
types, two males; one a pinned specimen, the other in alcohol, in the Cornell 
University collection. 

These specimens are all from Estes Park, Colo., August 7, 1921. 

♦Acknowledgments are due to Dr. O. A. Johannsen, who recognized this as a new species, 
and without whose assistance the following descriptions could not have been adequately made. 



398 The University Science Bulletin. 

LARVA. 

Length, 6-7 mm. when fully grown. Color white. 

Head very small, not over .25 mm. in diameter. Prothoracic proleg double, 
short, with many short spines; caudal prolegs short, each with a ring of about 
24 short spines. Anal gills not apparent. 

Head yellowish brown, with a narrow black hind border; on the ventral 
surface of the head, near the hind border, there is on each side a light oval 
spot, suggestive of an ocellus. Mouth parts very small and difficult to dis- 
sect out; t-hey are much reduced in size; the mandibles are composed of a 
single sharp tooth with two short spines on the inner margin near the base 
of the tooth; labrum small, the front margin excavated in the middle so as 
to leave a blunth' rounded chitinized tooth on each side; labial palpi very 
short : antennae minute, easily overlooked. 

PUPA. 

Length, 5-5.3 mm. Color brown; a narrow, blackish margin around the 
wings, and a narrow longitudinal dark line each side of the abdomen. The 
-^ntire pupa is smooth and devoid of any vestiture. Breathing trumpets not 
well developed, but represented by a spiracularlike chitinized area each side 
of the mesothorax. These breathing trumpets are best seen in the late 
larval stage when the pupa is being formed (fig. 7). Segment one to three 
of the abdomen smooth; segments four and five, above, each with a double 
transverse band of fine spines on the posterior margin, the spines of the hind- 
most bands directed forward and at least twice as long as the spines on the 
band immediately in front ; segments six, seven and eight) each with a single 
transverse band of short spines. Genital sacs smooth, no hairs or spines 
present. On the lateral margin of the abdominal segments the vestigial 
spiracles are visible. 

THE BIOLOGICAL RELATIONSHIP 

Between Trissocladms equitans and Rithrogena sp. 

With plentiful material on hand it has been possible to establish 
the relationship which exists between the immature stages of these 
insects. The larva of Trissocladius equitans is parastic upon the 
Rithrogena May-fly nymph. Although it has not been possible to 
learn in what manner the larva establishes itself as a parasite on the 
nymph, indications are that Trissocladius equitans spends its entire 
larval life as a parasite upon the May-fly nymph. Different-sized 
larvae, representing the instars, except possibly the first, were found 
under the wing pads of the nymphs. 

The parasite attaches itself to the posterior margin of the meso- 
thorax underneath the wing pads, where it imbeds its head in the 
softer tissue of the nymph and where it draws its nourishment from 
the host. The larva spins a sheet of silk, which completely invests 
its body, and by means of which it attaches itself firmly to the 
body of the nymph. At first, stretched to its full length, the larva 



Claassen: Larva of a Cpilronomid. 399 

lies across the body of the nymph, but as it grows hxrger the 
posterior end of (ho body doubles under, and later, when the larva 
becomes full grown, the middle part of its body projects backward 
over the abdomen of the nymph in the form of a letter U. When 
ready to pupate the larva releases its hold at the head end and turns 
back over the abdomen of the nymph. The pupa thus extends over 
about half the abdomen of the host. Tlic transparent sheet of silk 
invests the pupa as well as the larva. Just before the adult is 
ready to emerge the pupa breaks through this investing membrane 
and rises to the surface of the water, where the adult emerges in a 
manner similar to that in other chironomids. 

That the larva is a parasite upon the nymph is apparent from 
the following facts: First, the silk membrane completely sur- 
rounds the larva. There is no opening in this membrane which 
would make it possible to obtain food from the outside. Second, 
an examination of the stomach contents of a number of larvae 
failed to reveal the presence of any vegetable material, but did con- 
tain animal matter, especially fat bodies. This fact was also borne 
out in a study of cross sections of the larvse. Third, a study of 
the head and mouth parts of the parasite reveal their reduced con- 
dition. Tlie head is very small in proportion to the size of the body 
and the mouth parts are much reduced. 

The European species of the genus Trissodadius, of which the 
larvae are known, are all found as free living forms feeding upon 
vegetable matter.* 

The reduced size of the head and mouthparts of Trissodadius 
equitans indicate that this parasitic relationship has existed for a 
long time. Whether the parasite ever becomes so injurious as to 
kill its host the writer has not been able to determine. Another 
question of interest is what happens to the parasite when the nymph 
casts its skin. The life cycle of the parasite is either so short that 
development is completed during a nymphal instar of the host, or 
else the parasite must be able to detach itself from the cast skin 
and reestablish itself upon the newly emerged nymph or upon an- 
other individual, otherwise it must perish. 

There appears to be no relationship between the relative ages of 
the host and parasite. Pupa? of the parasite were found upon 
medium-sized nymphs, while, on the other hand, young larvse oc- 
curred upon nymphs that were evidently in the last nymphal in- 
star. 



*Barnard, K. H., in the Entomologist's Monthly Magazine, vol. 47:76-78, 1911, reports 
the case of a chironomid larva (name not given) which is parasitic upon a fresh water snail, 
Limnoea peregra. 



400 The University Science Bulletin. 



PLATE XLI. 

Fig. 1. Rithrogena nymph with a larva of Trissocladius equitans under the 
wing pads. 

Fig. 2. Nymph with a pupa. 



Claassen: Larva of a Chironomid. 



401 



PLATE XLL 





402 The University Science Bulletin. 



PLATE XLII. 

Fig. 3. Side view of May-fly nymph, showing the chironomid pupa under 
the wing pads. 

Fig. 4. Young larva of T. equitans under the wing pads of the nymph. 
Fig. 5. Full-grown larva on the nymph. 
Fig. 6. Pupa on the nymph. 



PLATE XIJI. 






^ fe- 



itii 

I 




A 




(403) 



PLATE XLIII. 

Fig. 7. Full-grown larva of T. equitans. In the thorax may be seen the 
breathing trumpets of the developing pupa within. 

Fig. 8. Mandible of the larva of T. equitans. 

Fig. 9. Labium of the larva. 

Fig. 10. Hind proleg of the larva. 

Fig. 11. Portion of tlie abdomen of the pupa. 

Fig. 12. Adult, T. equitans. 

Fig. 13. Clasper of male. 

Fig. 14. Wing of male. 

Fig. 15. Young larva. 

Fig. 16. Nearly full-grown larva. The black line indicates the edge of the 
silk membrane which envelopes the lana and by means of which it is 
attached to its host. 

(404) 



Claassen: Larva of a Chiroxomid. 



405 



PLATE XLIIL 




THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 17— October, 1922. 

(Whole Series, Vol. XXIV, No. 17.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

Water Insects from a Portion of the Southern Utah 

Desert R. C. Moore and H. B. Hungerjord. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE, KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 17. 



Water Insects from a Portion of the Southern Utah 

Desert. 

BY R. C. MOORE, 
Professor of Geology, University of Kansas ; and 

H. B. HUNGERFORD, 
Professor of Entomology, University of Kansas. 

INTRODUCTION. 
THE COLORADO PLATEAU. 

Not the least interesting of that well-known and yet little known 
country of varied attractions, the Great Western Cordillera of 
America, is the region of lofty plateaus, towering cliffs and deep, 
impassable canyons which is known as the Colorado plateau. 
Bordered on the east by the snow-clad peaks of the Rocky Moun- 
tains, on the north by the Uinta mountains, and on the west and 
south by low-lying deserts of the Great Basin and the lower Colo- 
rado valley, the Colorado plateau includes most of western Colorado, 
eastern and southern Utah, northern Arizona and northwestern 
New Mexico. Unlike the serrated peaks, irregular jagged spurs and 
sharp-topped divides of the Rockies', or of the mountain ranges in 
the Great Basin and Arizona deserts, the plateau country is a land 
of elevated, essentially flat-topped tables, which are terminated for 
the most part in steep, irregularly trending cliffs, and of great 
canyons which, converging on and culminating in the world-famous 
canyon of Colorado river, ramify almost every section of the plateau 
province. The tablelands are formed by hard rock formations 
which lie in more or less nearly horizontal positions, and the steep 
cliffs which border the plateaus or wall in the canyons mark the 
edges of these hard formations. Exceptions to the general archi- 
tectural scheme of the Colorado plateau country are a few small 
mountain masses of igneous origin, volcanic cones like the San Fran- 

(409) 



410 The University Science Bulletin. 

cisco mountains in Arizona, or laccolithic intrusions like the Henry 
mountains in southern Utah. These are not important in the aggre- 
gate, but are striking on account of contrast. 

All of the Colorado plateau region is arid or semiarid. There is 
little rain, and most of that which does come falls during a small 
part of the year and in torrential showers. Much of the character- 
istic topography and the aspect of the country in general is due 
chiefly to this. 

THE HIGH PLATEAUS OF SOUTHERN UTAH. 

The features and the conditions which are broadly typical of the 
Colorado plateau as a whole find especially characteristic expression, 
and, indeed, culminate in the south central part of Utah and adja- 
cent portion of Arizona. From the Grand canyon of Colorado river, 
the most profound and prodigious of the canyons, the stratified 
rock platforms rise tier on tier like gigantic stairs ascending north- 
ward. The top "stairs," in southern Utah, have an elevation of 
more than 10,000 feet above sea level, and comprise the so-called 
high plateaus. It is almost inevitable that this lofty plateau 
country, so closely adjacent to the deepest of the canyons, should 
be intricately dissected by tributary canyons. The high plateaus 
and adjoining region may therefore be specially designated as the 
"canyon lands." Travel is by tortuous and extremely toilsome 
routes, now in the depths of a profound abyss, now crossing a pla- 
teau spur of mountainous proportions. Some areas are absolutely 
inaccessible, and large districts are almost unexplored. Population 
is confined to a very few of the accessible valleys where irrigation 
permits cultivation of a little of the land adjacent to the water 
supply. Aside from these outposts of determined agricultural skill 
and industry, the region is traversed only by occasional cattlemen, 
prospectors or explorers. 

CLIMATE. 

The two main features which characterize the climate of southern 
Utah and affect more or less directly the life of the region, as well as 
physiographic processes, are dryness and temperature variation. 
In part of the area rainfall is less than five inches in the year, and in 
all of it the average is less than ten inches per year. In general, 
there is most rain in the three summer months, July August and 
September; and least rain in the spring months, April, May and June. 
The fall and winter months have an intermediate rainfall. The 
effect of the dry spring season, when most plants begin to develop 
rapidly and when many of the animals, especially the insects, pro- 



jNIoori: and Hiwckkford: Water Insects. 411 

gress swiftly through early life stages toward the vigor of mid- 
summer maturity, is to retard or to inhibit the development of life. 
Seeds may germinate, but, without rains, growth lags or ceases. 
Larva) of insects, polliwogs, and whatever animal life is dependent 
on waters, are restricted to permanent pools, springs or streams, for 
there are no temporary breeding waters in the dryness of the spring. 
The distribution of the yearly precipitation at four stations in the 
vicinity of the localities described in this paper, is shown in chart 
on page 412. 

The temperature of the plateaus is influenced to a large extent 
by the clear, dry atmosphere and by the altitude. During the days 
the sun warms the air and the ground, and the bare rocks reflect 
the heat rays back into the air. At night radiation is rapid and the 
temperature quickly falls. This large diurnal range in temperature 
is characteristic of arid or semiarid regions. At Tropic, in central 
western Garfield county, a freezing temperature in each of the 
twelve months has been reported, while the maximum temperature 
is greater than 100 degrees. (See chart, page 413.) 



412 



The University Science Bulletin. 



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414 The University Science Bulletin. ^ . 

surface "waters. 

Though the topography of southern Utah is very evidently the 
result chiefly of erosion by running water, there are few perennial 
streams in the region. The Colorado, master stream of the plateau 
country, gathers the waters from the west slopes of the Rockies, 
from the Uintas and other ranges where rains and melting snows 
furnish a varying but unending water supply. The tributaries 
which unite to form the Colorado begin, for the most part, as clear 
mountain streams, but in southern Utah the river carries so heavy 
a load of mud and sand that the water is dark reddish brown. On 
this account the name Colorado was first applied. The river is 
large and swift, with swirling eddies and numerous rapids. The 
only permanent tributaries to the Colorado in southern Utah are 
San Juan river on the south and Dirty Devil and Escalante rivers 
on the north. Each of these streams flows in a deep canyon, and, 
except for size, is essentially similar to the Colorado. The Esca- 
lante, which is the main stream in the region from which most of 
the collections of insects were obtained, is fed by the melting snows, 
rains and springs of the southern extremity of the highest of the 
plateaus, the Aquarius. Its waters are somewhat clearer than those 
of the other streams, because most of its course lies through massive 
sandstones. 

The numerous smaller streams in southern Utah contain water 
only at times of local rainfall. The run-off is very rapid, and after 
a torrential, muddy flood of less than an hour, or at most a few 
hours, the flow ceases. This type of swift, intermittent streams is 
characteristic of all the plateau region. 

After rains there are in places pools or "tanks" in such natural 
hollows as may retain any of the water. Some of these are found 
in the stream channels, others in eroded depressions in the nakecf 
rock. Some are small, shallow and short-lived under the rays of 
an intense sun and a thirsty atmosphere; others are larger, and 
occasionally deep. Those in the channels of streams commonly 
contain very muddy water, which, because of the excessive fine- 
ness of the mud, retains the sediment in suspension until evapora- 
tion gradually gives it the consistency of thick soup or gravy, and 
finally of brick. Where the rain water accumulates in sandstone, 
or where the gathering waters do not cross exposures of the soft 
shales which furnish most of the muds, the waters remain fairly 
clear. The larger "tanks" may retain water from year to year, 
shrinking slowly through times of drought, but refilling on the 



Moore AND Hixgerford: AVatkk Insects. 415 

coming of rain. These little pools of moderately clear water afford 
sanctuary for such water-breeding or water-living animals as in- 
habit the region, and in some cases it is a populous and assorted 
community that crowds together. 

Springs may be mentioned among the surface waters of southern 
Utah, although they are not numerous, and the water from them 
very short h^ sinks into the ground. They are in many cases nearly 
permanent, and are an important source of supply, especially as 
regards the uses of man. None of the springs or seepages in the 
plateau region studied has a fiow of more than a few gallons an 
hour. 

It is freciuently a number of miles from one spring, ''tank" or 
other permanent water source to another. 

FIELD WORK. 

In the summer of 1921 and 1922, Mr. Moore, with a party of four 
assistants, was assigned by the United States Geological Survey to 
make an examination of a portion of the high plateaus of southern 
Utah, with reference to coal resources and possibilities of oil and 
gas development. A detailed geological map of approximately 
3,000 square miles was made, and reconnoissance study of a very 
much larger area was completed. The region mapped comprises 
most of central eastern Garfield county, a portion of central Wayne 
county, and eastern Kane county, Utah. Although the primary 
purpose of this work had to do with possible coal, oil and gas re- 
sources, special attention was given to water supply, for the region 
is a^emidesert. Not only was it important for the party to find 
water for camp purposes, but the possible development of water 
supply is in all cases the most important consideration in the utiliza- 
tion of such a region. During the course of this work, wherever 
water insects were found, and where it was possible without inter- 
ruption of the main objects of the work, collections were made. 
Since no other similar collections have been reported from this 
region, and since the distribution of water insects in this region is 
both of biological and general entomological interest, the results 
are here presented. Identification and special notes on the species 
found are the work of Mr. Hungerford. 

DESCRIPTION OF COLLECTIONS. 

Since the collections of water insects to be noted below were 
gathered as opportunity offered and not as a part of a systematic 
faunal survey, the data are perhaps somewhat fragmentary and 
scattered. However, since the sources of water supply in such 



416 The University Science Bulletin. 

country are not numerous, and since it is necessary for the traveler, 
whatever his mission, to seek the places where water may be had, 
it is probable that a considerable number of the places where water 
insects might occur were observed. In each of these a collection, 
representing as far as possible all of the species present, was ob- 
tained. The collections include mainly those from springs and 
"tanks." It is possible that water insects could have been found 
in some of the seepages along streams or in the permanent streams, 
but there was no opportunity to make special search for these. 

SMALL "tank," one AND ONE-HALF MILES ABOVE MOUTH OF MULEY 
TWIST CREEK, EASTERN GARFIELD COUNTY, UTAH. 

Muley Twist creek is an intermittent stream whose bed is dry 
the greater part of the year. It occupies a very deep box canyon, 
carved in massive red and yellow sandstone. About seventy-five 
feet above the bottom of the canyon, on its sloping west sandstone 
wall, was found at one point about one and one-half miles above 
the mouth of the creek, a little "tank" about one by two feet in 
width and length and with greatest depth of approximately one 
foot. The depression was filled with clear water, part probably 
caught in a recent rain and part derived from a very small seep in 
the sandstone above. The "tank" is probably not at all permanent. 
It was surrounded by bare smooth sandstone, without any near-by 
plant growth. 

In this little basin lived a group of seven individuals of Arcto- 
corixa abdominalis Say, without other insect life. 

"tank" about two MILES ABOVE MOUTH OF MULEY TWIST cllEEK, 
EASTERN GARFIELD COUNTY, UTAH. 

On the east side of the Circle Cliffs and along the Water Pocket 
Fold in eastern Garfield county are a number of "tanks" which con- 
tain water in all but the driest seasons. From one of these, in the 
bed of Muley Twist creek, beneath a high sandstone cliff, a number 
of water bugs were taken. The pool was about eight feet in 
diameter and three feet deep in one part at the time of our visit in 
August. Two or three large cottonwoods shade the pool, but there 
is almost no other vegetation near at hand. 

In this little basin was taken one dytiscid beetle, Rhanhis bino- 
tatus Harr., and upon the surface eight specimens of Gerris orba 
Stal, all winged, and one pair mating (five males and three females). 
The solitary dytiscid beetle appears to be a new record for Utah. 
Its previous capture has been recorded for Wisconsin and Arizona. 



Moore and Hungerford: Water Insects. 417 

"tank" on COLORADO RIVER AT WATER POCKET FOLD, KANE COUNTY. 

This was located on the north bank of the Colorado river near the 
mouth of a small tributary canyon at Bennett's oil camp, where 
Water Pocket Fold crosses the canyon of the Colorado river, about 
eight miles above the mouth of Escalante river, eastern Kane 
county. A pool of clear rain water filled the depression in bare 
sandstone, about four feet wide, five feet long and two and one-half 
feet deep; no vegetation around the pool; about thirty yards from 
Colorado river, a swift, muddy stream, very unlike the water in the 
"tank"; unknown distance to adjacent pools of clear quiet water; 
elevation about 3,500 feet. Here were taken: 

Fifteen Arctocorixa abdominalis Say, four males, eight females 
and three nymphs. This represents the farthest north record for 
the species, which was described from Mexico and reported from 
Texas and California by Uhler. 

One Notonecta insulata Kirby, female. 

Two Thermonectes marmoratus Hope, one male and one female. 
The capture of this beautiful dytiscid appears to be a new record for 
the state. Leng's catalogue records it from Arizona and Lower 
California. 

SPRING OPPOSITE OWL CAVE, HARRIS WASH, GARFIELD COUNTY, UTAH. 

About twenty miles southeast of the little town of Escalante, 
eastern Garfield county, Utah, a permanent spring is found in the 
sandstone wall of Harris canyon, which is tributary to Escalante 
river. The fiow of this spring is not large, but the water is of excel- 
lent quality, and, as natural shelter is conveniently near at hand in 
a large sandstone caveYn, Owl Cave, the spot is a camping place for 
most of the few travelers who pass this way. The water from the 
spring irrigates naturally several square yards of protected ground 
adjacent to the spring. A beautiful little meadow, therefore, has 
been formed, through which the water flows a short distance before 
sinking into the sand. 

This spring and short stream support a populous community of 
aquatic insects and other life. However, only two species of water 
bugs were observed, Gerris remigis Say and Trepobates pictus Uhl. 

SPRING IN WATER POCKET FOLD, CIRCLE CLIFFS, GARFIELD COUNTY, 

UTAH. 

One of the very few water sources in the northeastern side of the 
Circle Cliffs is a small spring, from which the water flows a few 
yards before disappearing in the floor of a canyon tributary to 



418 The University Science Bulletin. 

Muley Twist creek. The adjacent rocks, chiefly sandstone, are 
nearly bare of vegetation, though not far distant are numerous 
pinon, scrub cedar and sagebrush. The spring varies somewhat in 
volume, but does not commonly dry up in the spring dry season. 

On the surface of the spring and the little stream below are 
numerous representatives of the species Trepobates pictus and Gerris 
remigis. A large number of individuals of each were obtained, in- 
cluding a considerable number of nymphs belonging to the first 
species. The Trepohates were more numerous than the Gerrids at 
the time of obser^'ation in August, 1921 — a rather unusual condition 
as compared with other localities inhabited by the two forms. 

SPRING on upper HENRIEVILLE CREEK, SOUTH CENTRAL GARFIELD 

COUNTY, UTAH. 

An area of special scenic and geologic interest in southern Utah 
is Table Cliff and the surrounding very rough district in south 
central Garfield county. South of Table Cliff, on one of the branches 
of upper Henrieville creek, is a seepage of very alkaline water, the 
only source available for camp use in the head canyons under the 
cliff. The water is so strongly mineralized as to be hardly potable. 

On the surface of the short, trickling stream below the seep several 
water bugs were taken. A few Gerris remigis Say and a considerable 
number of Micro velia are the inhabitants of this apparently in- 
hospitable haven. The Microvelia species of this region has not 
been determined. It may be new. 

"tank" NEAR SOUTH POINT OF KAIPAROWITS PLATEAU, EASTERN 

KANE COUNTY, UTAH. 

Most of the observed watering places in southeastern Utah did not 
contain water insects at the time of visit in the summer of 1922. Of 
four springs and pools on the top of Kaiparowits plateau, a high 
tableland which extends southeastward toward Colorado river from 
the uplands farther north, only one appeared to contain any 
aquatics. This was a partially rain-filled "tank" in the bottom of 
one of the narrow, steep-sided canyons which cut the plateau sur- 
face. The "tank" was carved in bare, massive sandstone, without 
near-by vegetation. The pool was about six feet long by one foot 
wide, and its maximum depth a little over one foot. In this pool 
were several Notonecta insulata Kirby, but no other water bugs. 



MooHK AM) Ih xgerford: Water Insects. 419 

LAST CHANCE CREEK, EASTERN KANE COUNTY, UTAH. 

In ;i pnrt of the sandstone canyon of Last Chance, about twenty- 
five miles :il)()\-e the point where it flows into Colorado river, a 
small stream Hows more or less permanently. Other parts of the 
canyon contain intermittent flow, but in most cases, except for 
occasional groups of Gerris remigis, did not contain other observed 
water bugs. At the place described, a group of small dytiscids, a 
number of MicrovcUa and some gerrids were collected. 

SPRING ON PARIA RIVER, ABOUT FIFTEEN MILES BELOW CANNONVILLE, 

KANE COUNTY, UTAH. 

Most of the lower Paria river, the main stream in south central 
Kane county, Utah, is dry the greater part of the year. The river 
has carved a very deep sandstone canyon, which affords the only 
passageway from north to south across this part of the country. At 
a point about fifteen miles below the town of Cannonville, the last 
settlement on the southward route to Colorado river, at Lee's Ferry, 
about seventy-five miles distant, is a fine spring. The flow is fairly 
large, as compared with other springs in this country, and the water 
is clear and not alkaline. 

A host of Gerris remigis and Trepobates pictus were found on the 
water of the spring. 

LOWER ROCK CREEK, EASTERN KANE COUNTY, UTAH. 

Water fl.ows more or less permanently in the lower part of Rock 
creek, the easternmost of the longer, deep canyons tributary to Colo- 
rado river west of Kaiparowits plateau. The canyon of Rock 
creek, as the name perhaps suggests, is somewhat unusual, even 
among the great, bare rock cliffs and canyons of the plateau coun- 
try. Its walls and bottom are mostly composed of naked rock, and 
sculpture by running water has produced an almost indescribably 
rough topography. 

The stream at the camp site of the writer, about two miles above 
Colorado river, contained numerous Microvelia and a few gerrids. 
No other water bugs were observed. 

NEAR GOODRIDGE, SAN JUAN COUNTY, UTAH. 

In a small, isolated spring, about one foot in diameter and a few 
inches deep, were taken three Notonecta insulata Kirby. No other 
water insects were present. This tiny basin of water was three miles 
from the swift, muddy San Juan river, and there were no other near 
water sources known. The location of tiie spot is at the west side 



420 The University Science Bulletin. 

of Comb Ridge, where the road from Bluff to Goodridge enters 
Comb Wash. Some vegetation was growing about the spring. 

To the above Utah collections may be added this interesting one 
from northwestern Colorado, made in 1920. 

junction mountain, YAMPA river, northwestern COLORADO. 

In the deep canyon of Yampa river, where the river plunges 
through Junction mountain, about thirty-five miles east of the con- 
fluence of the Yampa and the Green, in northwestern Colorado, were 
taken forty-four Rhagovelia distincta Champ., all apterous; nine 
were males, four nymphs, and the remainder females; also seven 
Trepobatopsis trux Bueno, two males and five females, described by 
Mr. Bueno as new. 

From a small, sluggish stream at west side of Junction mountain 
were taken an interesting series of large gerrids. These vary in 
color from russet to the dark color typical of Gerris remigis Say. 
The russet-colored form has been determined by Mr. J. R. de la 
Torre Bueno as Gerris orba Stal. Of these there are nineteen speci- 
mens, nine males and ten females. Thirteen of the lot are apterous. 
G. orba Stal has been listed from California, Oregon and Nevada 
hitherto. The others, forty-seven of them, all apterous, are of a 
trifle lighter color than G. remigis Say, but obviously are the same 
as Bueno's orba. 

In an attempt to find differences between our common Gerris re- 
migis Say and this series from the West, all of the Kansas and Colo- 
rado material available (a series from the eastern margin of Kan- 
sas through the state and across Colorado) has been very carefully 
studied. The Kansas forms are darker and larger as a series, but it 
does not seem possible at the present time to fix upon any structural 
characters to distinguish these forms. 

In connection with the problem of separating the striders, it may 
not be out of place to state that it seems to have been overlooked 
that Gerris conformis Uhl. males have the ventral side of the sixth 
abdominal segment singly emarginate, like Limnoporus rufoscutel- 
latus Latr. Indeed, the two species have, in addition to the above 
genital character, another distinctive character that is common to 
both, namely, their very long legs. The hind femora in both spe- 
cies greatly surpass the tip of the abdomen. The examination of 
such material as has been available suggests that the two species 
commonly accepted as L. rujoscutellatus and Gerris conformis Uhl. 
may be separated as follows : 



Moore AND HuNGERFORD : Water Insects. (^y ^^/ 

A. Antennal segment 1 longer than 2 plus 3. Eyes protuberant. Middle and 
hind femora about equal in length. Venter of sixth abdominal segment 
in male with median longitudinal fossa. The seventh with a longitudinal 
ventral carina. Gerris conjormis Uhl. 

AA. Antennal segment 1 shorter than 2 plus 3. Eyes not protuberant. Middle 
femora shorter than hind femora. Venter of sixth abdominal segment in 
male without fossa and seventh segment not carinate. 

Liinnoporus rufoscutellatus Latr. 

In addition it may be stated that L. rufoscutellatus Latr. is a 
more slender species, and that it is usually of a russet color. Gerris 
conjormis Uhl. has the anterior part of the pronotum bearing two 
prominent papillae and well-elevated callosities. 

It would seem somewhat of a question whether these two species, 
having in common genital characters and exceptionally long femora, 
can be placed in separate genera upon the antennal character alone. 
Structurally they are more nearly congeneric than G. remigis Say 
and G. conjormis Uhl. The latter belongs with L. rujoscutellatus 
Latr., and the generic characters of Limnoporiis should be revised. 

resume. 

In brief resume of the above collecting notes, it may be said that 
the southern Utah desert affords extremely interesting problems for 
the student of water life. The light' annual rainfall, the isolation of 
the pools and the nature of the water they contain, lend interest to 
the study of the resident population of the waters. The collections 
here reported represent the complete insect population of the various 
pools surveyed. One isolated little spring-fed pool, the size of a 
washbasin, contained three Notonecta insulata Kirby, and no other 
insects. Another little rain-filled "tank," less than two feet in diam- 
eter and a foot deep, located high up on the sloping wall of a canyon, 
contained seven water boatmen, Arctocorixa abdominalis Say, and 
not another insect. The scant annual rainfall of from five to ten 
inches, with the exceptionally light fall in April, May and June, 
works against a large population of aquatics. The water bugs 
thus utilize every available supply of water. The pools contained 
the corixids, notonectids and dytiscids, while the striders were found 
on the springs more commonly than upon the "tanks." The pres- 
ence of winged species in places remote from permanent water is 
readily understood. The finding of eight Gerris orba Stal, all 
winged, upon a rain-filled "tank" and forty-seven specimens of the 
same species along the Yampa river, all apterous, is worthy of note.* 

* An interesting note on the factor regulating wing development is suggested by collect- 
ing notes on Microvelia macgregori Kirk, in New Zealand by Mr. J. G. Myers, in the New 
Zealand Journal of Science and Technology, vol. V, pp. 6 and 7. He writes that his col- 
lections of this insect in the lowlands have been almost entirely apterous forms, while at an 
altitude of 4,500 feet every capture was winged. 



THE 

KANSAS UNIVERSITY 

Science Bulletin 

Vol. XIV, No. 18— October, 1922. 

(Whole Series, Vol. XXIV, No. 18.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

The Xepid.e in North America North of Mexico, 

H. B. Hungerford. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE KAN. 



Entered at the po.st office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 18. 



I 



The Nepida3 of North America. 

(Further Studies in Aquatic Hemiptera.) 

BY H. B. HUNGERFORD. 

ACKNOWLEDGEMENTS. 

WISH herewith to acknowledge my indebtedness to those who 
have rendered assistance in the preparation of this paper, which 
had its beginning several years ago. I am under especial obligation 
to Dr. L. O. Howard, Dr. J. M. Aldrich, and Mr. W. L. McAtee for 
the opportunity they gave me to study in the United States National 
Museum, where are located some of the types of Doctor Montandon 
and the Kirkaldy and Uhler collections of these insects. In addi- 
tion to the material at the United States National Museum, I have 
been permitted to examine the collections of Mr. J. R. de la Torre 
Bueno, of Dr. Carl J. Drake and have had, beside our own Univer- 
sity collections, abundant material so generously supplied by Mr. 
W. E. Hoffmann and Doctor Knight, of the University of Minne- 
sota, and by Mrs. Grace Wiley from collections in Texas. Profes- 
sor Brimley, of Raleigh, N. C, also supplied me with two species 
for study. Miss Kathleen Doering made most of the drawings and 
Mr. P. A. Readio and Mr. Raymond Beamer helped me with the 
photographs. 

INTRODUCTION. 

The subtle characters and elusive specific differences between the 
various species of the cryptocerate groups of the Hemiptera have 
made it difficult to fix specific limits. The students of these groups 
have resorted, therefore, to comparisons that are beyond the compre- 
hension of the general systematist endeavoring to identify his col- 
lection. Such comparative notes become intelligible only after a 
prolonged study of the insect group concerned, and this condition 

(425) 



426 The University Science Bulletin. 

has worked against there being a very large number of students 
who have found the time and inclination to identify the species in 
such families as the Notonectidse, Nepida? and Corixidse, for in- 
stance. 

The difhculty encountered by the writer in sorting and naming 
the aciuatic bugs in the course of his biological studies led to a thor- 
ough investigation for diagnostic characters of more demonstrable 
kind than those employed in the literature. It has been the en- 
deavor, therefore, to find and figure characters of sucli definiteness 
that anyone with even fair training in close work can be certain of 
the species he is trying to identify. 

The first report upon this work was given under the title, "The 
Male Genitalia as Characters of Specific Value in Certain Crypto- 
cerata," which appeared in volume XI of the Kansas University 
Science Bulletin, December, 1919. In this the writer called at- 
tention to these characters in the Naucoridse, Saldidae (not a crypto- 
cerate, of course), Gelastocoridae, Corixidse and Notonectidse. The 
treatment of the genus Notonecta was sufficiently complete to be of 
value to the systematist, because the male genital capsules of all the 
North American species were figured or described, save two, A\ iM- 
eri Kirk, and N. montezuma Kirk., the former readily recognizable 
by the description, and the latter exceedingly rare, repre.>:nted, so 
far as indicated in the literature, by two specimens in the Hope 
museum, carefully studied and figured by Champion. (Figure 8, 
plate XXXI, Science Bulletin XI, figures Notonecta howarclii Bueno, 
and fig. 5, plate XXXI, Science Bulletin XI, figures Notonecta 
shooteriiVhl. The one there named N. lutea is, of course, N. borealis 
Hussey and Bueno.) 

Studies were made upon the Nepidse and Corixidae, but were not 
reported in the first paper, and there was no intention, until recently, 
of publishing upon the Nepidse. However, four species of Ranatra 
have been taken in Kansas waters as a result of recent collecting, 
all new records for the state. The task of naming these water scor- 
pions involves problems in nomenclature and the authenticity of 
species — matters concerning which there is a difference of opinion 
among our best authorities. 

Mr. J. R. de la Torre Bueno has believed that Ranatra nigra Her- 
rich Schaffer 1853, is a synonym of Ranatra fusca Palisot Beauvois 
1805. The insect which he calls Ranatra fmca is large with very 
prominent eyes, slender anterior legs, and a deep, broad prosternal 
groove — a very characteristic insect. With this interpretation of 



Hlxgekfurd; Xki'ii).!-: im Amerra. 427 

Palisofs species, he described a small, compactly built insect with 
\ei'v broad front femora, as Ranatra kirkaldyi. 

Dr. A. L. Montandon (Bui. Soc. Sci. Bucharest, XIX, 1910) con- 
siders Ranatra kirkaldyi Bueno a synonym of Ranatra fusca P. B., 
and concludes that Ranatra nigra H. S. is a good species, although 
he has seen no American specimen that fits the description. Van 
Duzee follows Montandon. 

In a recent issue of the Entomological News (vol. XXXII, p. 
273, 1921), under the title "New Records of Aquatic Hemiptera for 
the United States with Description of New Species," Mr. Torre 
Bueno again defends his Ranatra kirkaldyi and comments on Ra- 
luitra fusca P. B., describing it so closely that there can be no mis- 
taking the insect, and stating that there is a specimen of this species 
in the United States National Museum labeled Ranatra fusca P. B., 
by Doctor Montandon. This statement is indeed true. 

Through the kindness of Dr. L. O. Howard, Doctor Aldrich and 
W. L. McAtee; the writer had the pleasure of spending some time at 
the United States National Museum studying the aquatic Hemip- 
tera. He had the opportunity, therefore, of examining the material 
determined by Doctor Montandon, as well as the types of Ranatra 
protema Montd., Ranatra brevicollis Montd. and Curicta hoivai'dii 
Montd. He found two distinct species identified by Doctor Mon- 
tandon as R. fusca P. B. One of them is Bueno's Ranatra fusca; 
the other, cjuite a different species, but not Bueno's R. kirkaldyi. 

After examining much material in the family Nepidse, and very 
carefully studying and weighing original descriptions and compara- 
tive notes, the writer has been forced, in spite of his reluctance, to 
a new interpretation of the old species. The facts which led to the 
change in nomenclature are presented under the species concerned. 
All the species are figured in such detail that there need be no ques- 
tion or uncertainty as to designation, and these studies should aid 
in arriving at a solution of the matters in controversy. 

Tlie family Nepidse is represented in America, north of Mexico, 
by three genera, Nepa, Curicta and Ranatra, which may be sepa- 
rated as follows: 

A. Body broadly oval and flat. Nepa. 

AA. Body elongate. 

B. Prothorax a little broader than head, body elongate oval. 

Curicta, 

BB. Prothorax narrower than head, body ver>^ elongate. Ranatra. 

Genus Sepa, see p. 428; genus Curicata, see p. 429; genus Ranatra. see p. 433. 



428 The University Science Bulletin. 

Genus Nepa Linnaeus 1758. 
We have one species in the genus Nepa: N. apiculata Uhl. 

Nepa apiculata Uhler. 

Uhler, P. R., in T. W. Harris' Insects Injurious to Vegetation, 3d ed., p. 12, plate 1, fig. 
1 f 1862. 

This species was first made known to science through a picture of 
it which appeared in the third edition of Harris' "Insects Injurious 
to Vegetation." In this edition Doctor Uhler added notes on the 
Hemiptera. Then in 1878, Uhler, in his "Notices of the Hemiptera 
Heteroptera in the Collection of the Late T. W. Harris" (Proceed- 
ings of the Boston Society of Natural History, vol. XIX, Pt. IV, 
1878), records the presence of "No. 26, Harris Collection, Nepa apic- 
ulata Say MS., under stones near water, May 15, 1826," and adds 
the following descriptive note: "The principal differences between 
our species and the European one consist in the color of the tergum, 
which is red in the latter, fuscous in ours ; and the length of the api- 
cal tubes, which in ours are stouter and shorter." In commenting 
on these comparisons, Montandon (Bui. §oc. Sci. Bucharest, VIII, 
1898) says that Uhler, like Stal and Ferrari, attached too much im- 
portance to color, and gives illustrations of the variability of Nepa 
apiculata Uhl. and of Nepa cinerea L. to prove the danger of con- 
sidering color of specific significance. He then states that the Amer- 
ican species is a little more transverse across the thorax, the abdo- 
men proportionately larger in the rear, and the respiratory tubes 
shorter than in the European species. Again in volume XVIII of 
the same periodical, under the title, "Hydrocorises de I'Ame'rique du 
Nord, Notes et Descriptions d'Especes Nouvelles," Doctor Mon- 
tandon adds that in "Nepa apiculata Harris 1862, Uhler 1847,* 
. . . the superior part of the head is less boldly carinate, espe- 
cially on the vertex, which is generally almost smooth, cjuitc feebly 
convex." 

These differences between the European Nepa cinerea L. and the 
American Nepa apiculata Uhl. do exist, as an examination of the 
photographs on plate LI will show. However, without both species 
for study, one would be compelled to remain in doubt or name the 
species from its geographical distribution. Since there do occur 
marked structural differences, it is well to note them. The antennae, 
for instance, are very different, as an examination of figures 5 and 

*Probably a typographical error, because Uhler was born in 1835. Van Duzee omits the 
reference: Montandon, Bui. Soc. Sci. Bucharest, XVIII, p. 180, from the list under jVepa 
apiciblata Uhl. in his catalogue of 1917. It is interesting to note that Nepa apiculata was a 
Say manuscript species. 



Hungerford: Nepid^ in America. 429 

6, on plate XLVII, will show. The penultimate segment of the 
European form has a lateral prolongation which gives the antenna 
a branched appearance. The Nepa apiculafa lacks this entirely. 
The male genital capsules are also unlike. The considerable differ- 
ence between these and between the claspers is indicated on plate 
XLVII, figures 1, 2, 3 and 4. 

The nearest approach to a formal description of the American 
species is given by Uhler in the Riverside Natural History, vol. II, 
p. 253, 1884. It follows: 

Color (lull fuscous gray, with the base of the abdomen above more or less 
tinged with reddish. It is of an elliptical form, blunt in front, with a ridged 
middle line on the vertex, and with three short raised lines on the prothorax. 
each side of a longer one on the middle. The surface and margins of the 
thorax and head are roughly granulated, while these, tog.ether with the scu- 
tellum and corium, are rough and closely covered with stiff, short pile. The 
anterior femora have no teeth on the inner angle, but instead there is a prom- 
inent elbow, forming a wide expansion for the sides of the deep gutter. The 
wings are smoke brown, with darker veins. This species closely resembles the 
European one, and measures about twc^-thirds of an inch to the end of the 
abdomen; while the respiratory tubes are a little more than one-fourth of an 
inch in length. 

Montandon has shown the differences between our species and 
the European A''. Cinerea Linn. 

The United States National Museum has specimens from Massa- 
chusetts, Pennsylvania, Maryland and Illinois. I have seen speci- 
mens from New York, Minnesota and Wisconsin. 

Genus Curicta Stal, 1861. 
These insects are very interesting because they are intermediate 
between the broad, flat Nepa and the very slender Ranatra. The 
Americas can boast of several species in this genus, but for North 
America, north of Mexico, there has previously been recorded only 
one, and of it but a single specimen, taken at Victoria, Tex., just 
within our limits. From this specimen Doctor Montandon drew up 
the description of Curicta howardii. It is a pleasure, therefore, to 
record the capture of other specimens of this species and to add a 
second species which has not been described. This second species 
has been the subject of life-history studies by Mrs. Grace Wiley, 
and is reported further on in this bulletin. 

KEY TO SPECIES. 

A. Lateral prolongation of penultimate segment of antenna very short (see 
fig. 8, pi. XLVII). C. howardii Montd. 

AA. Lateral prolongation of penultimate segment of antenna very long (see 
fig. 7, pi. XLVII). C. drakei sp. new. 



430 The University Science Bulletin. 

. Curicta howardii Montandon, 1910. 

Montandon, A. L., Bui. Soc. Sci. Bucharest, XVIII, p. 181; 1910. 

The original description is in French. The writer's free trans- 
lation follows: 

Elongate oval in form, visibly attenuate in front and rear, lateral margins 
not subparallel, the greatest width toward the posterior third. Head quite 
enlarged, although a little narrower than the front part of the pronotjuii, as 
long as wide, including the eyes, longitudinally carinate throughout its length, 
the carina more obtuse on the posterior interocular portion. Interocular 
space more than three times as wide as the diameter of the eye. Eyes small, 
globular, anterior part of head triangular, exceeding the anterior level of the 
eyes by a length ecjual to its width between the eyes in front. 

Pronotum distinctly longer than its width behind, lateral edges subparallel 
on their anterior three-fifths, quite strongly widened on their posterior two- 
fifths; with four obtuse longitudinal carinse, little accentuated and sub- 
parallel, two each side of the anterior part, the posterior part with two 
oblique carinae arising from the anterior median carina and quite divergent 
behind. The anterior depression ' of the pronotum broadly semicircular, the 
anterior angles quite narrowed, subacute. 

Scutellum with three longitudinal carinas, the median continuing quite 
plainly clear to the apex of the scutellum. The two laterals slightly diverging 
behind, ^•anishing on the middle of the sides of the scutellum, which are 
slightly sinuate before the tip or end. 

Coria insensibly and gradually widened behind on their basal halves, at- 
taining their greatest width behind the middle and narrowing thereafter; 
membrane well developed, regularly subrounded at the extremity. Com- 
missure of the clavus almost twice as long as the scutellum. 

Appendages short, quite robust toward the base, attenuated thereafter, 
about half the length of the abdomen. 

Anterior femora quite robust, as long as the pronotum on its lateral edgeSj 
with a single median tooth easily visible on the inner edge of the groove where 
the folded-up tibia is lodged, this tooth distinctly closer to the base than to 
the apex of the femur; the external side of the groove also appears denticu- 
late, as if notched on the basal third of the femur. Neither teeth nor sinuosi- 
ties toward the apex of the femur. 

Anterior coxae half the length of their femora. Anterior tibia quite long, 
blackish, with a pale annulation toward the base, and the apical third like- 
wise pale; the extremity of the tarsi come to the basal third of the femora 
when the tibia is folded back against the latter. 

Intermediate and posterior legs short, the ends of the posterior femora, 
which are a little shorter than their tibiae, do not reach the suture of the last 
abdominal segment. Intermediate and posterior tarsi with their claws less 
than half as long as their tibiae. 

Median longitudinal part of the prosternum slightly saddle-shaped, pro- 
jecting in all its width, more elevated than the lateral pieces, a little flat- 
tened and traversed its whole length by a fine median groove; very obtusely 
tuberculate in its anterior part. A greater space between the intermediate 
coxae than between the anterior or posterior coxae. 

Length, 19 mm.; maximum width a little behind the middle of the corium, 



Huxgkkfokd: Nkpid.?<: in America. 431 

4.5 mm.; at bn^^v ol' I'lonotum. 3.8 mm.; length of appendages, 7.7 mm., Vic- 
toria. Tox. A single specimen, United States National Museum, Washington. 

Tt) the above description Doctor Montandon adds the following 
helpful coniparative notes: 

This species is intermediate in size between C. volxemi Montd. and C. 
Scorpio Stal (.=>nonta)idom Martin). It differs from C. voh-cini Montd. by 
the iironotiun being sensibly narrowed in front, while in the latter it is almost 
as wide in front as behind. In this character it approaches more closely to C. 
Scorpio Stal, which also has the pronotum quite narrowed in front, with the 
same right-angled anterior angles almost sharp, but its anterior tibife are, 
however, slightly more elongated than in this latter species; that is to say, 
much more than in C. volxemi Montd. In the character of the median tooth 
of the anterior femur being situated closer to the base than the extremity, 
howe\-er. the species approaches C. scorpio Stal, but it is plainly separated 
from the latter by its head being longitudinally carinate throughout its entire 
length, while the head is simply convex between the eyes in C. scorpio Stal 
and almost plain in C. volxemi Montd. 

Furthermore, in C. scorpio the scutellum is not carinate; the longitudinal 
grooves of the pronotum are also much less emphasized and the anterior 
tibiae are more largelj* pale, darker only toward the base. 

This is the first species of the genus found in the United States. It is 
to b(> ]iresumed, however, that others may occur in the Southern states 
neighboring to Mexico, where are foimd the two species to which I have 
just compared it. 

I take pleasure in dedicating this to Mr. L. 0. Howard, as an indeed 
feeble expression of my sincere gratitude. 

In the collection of Prof. H. G. Barber are three males from 
Huachuca mountains, Arizona, taken in 1899. They have been 
compared with the type in the United States National Museum 
by Dr. Carl Drake, through the kindness of whom I have been 
privileged to study the structural details of these interesting in- 
sects. The United States National Museum has a specimen taken 
by D. C. Van Dine at Camp Travis, Tex., March 30, 1918. 

The species is a compactly built creature. The head is set 
deeply into the prothorax, the anterior lateral lobes of which are 
conspicuously swollen and incurved on their anterior tips to em- 
brace a portion of the eyes. The front of the head appears almost 
truncate, due to the tylus and juga being about equal. The 
antennae are small and the lateral prolongation of the penultimate 
segment short (see fig. 8, pi. XLVII). The pronotum has a broad, 
prominent elevation throughout its length. On the anterior half 
of this there is a median longitudinal depression. The genital 
capsule and its claspers are somewhat different from those of the 
species described below (see figs. 7 and 8, pi. XLV) , but the specific 
differences are not great. 



432 The University Science Bulletin. 

Curicta drakei sp. new. 

Size. Length from 16 mm., in case of smallest male, to 22 mm., in largest 
female — not counting the respiratory filaments, which are from 7 to 8 mm. 
long. The width of the head, including the eyes, from 2 mm. to 2.25 mm.; 
the greatest width of anterior part of thorax, 2.5 mm. to 2.8 mm.; the 
greatest width of posterior portion of thorax, 3 mm. to 4 mm.; the greatest 
width of abdomen, 3.6 mm. to 4.75 mm.; the length of pronotum measvu-ed 
along the dorsal median line, 3, mm. to 3.6 mm. 

Color. Obscured by incrustations, which color it from gray to black. 
The cleaned insect is yellowish to dark testaceous, the dorsum of abdomen 
red. 

Shape. Relatively slender; greatest width of thorax is to length of body 
approximately as 1 is to 5.4. Sides of thorax and abdomen nearly parallel. 

Structural characteristics. The head wider across the eyes than distance 
from caudal margin of head to tip of lora. The length before the eyes greater 
than that behind them. Ej^es small, globular. Tylus longer than juga. The 
head longitudinally carinate. 

Penultimate segment of antennae with long, slender lateral prolongation 
(see fig. 7, pi. XLVII). The pronotum with anterior lateral lobes not greatly 
swollen; the lateral margins, therefore, not greatl.y curved. Three longitudinal 
grooves on disc of pronotum, the median one broad and straight, dividing 
the median longitudinal elevation into two parallel carinse, laterad of which 
are the deep lunate grooves terminating caiidolaterally in deep depressions. 
The posterior enlarged portion of pronotum bears four more or less distinct 
longitudinal carinae, which in some specimens appear to arise as bifurcations 
of the two prominent carinse before them, and again as independent eleva- 
tions. Scutellum tricarinate, the median carina more prominent on posterior 
half, the lateral carinse slightly curved and terminating about the middle of 
the lateral margins of the scutellum, these elevations and declivities often 
accentuated or obscured by incrustations. Membrane of wings well de- 
veloped, as long as the elytral suture, reticulate, the apex of the wing cover- 
ing the basal two-fifths of the genital segment. Operculum of male genital 
segment semicylindrical and slightly constricted before the apex, which is 
bluntly pointed and faintly carinrtte. The metasternal plate short, caudal 
margin nearly straight, the posterolateral prolongations attaining less than 
half the length of the coxae and exposing a large elevated area (as long as 
wide) of sternite behind it. Front coxa and femiu- stout, coxa one-half length 
of femur, which bears its rather well-developed tooth nearer the base than the 
apex. This tooth is located on the inner edge, two-fifths of the distance from 
the trochanter to the apex of the femur. Tibia one-half as long as femur, 
the apex of the short tarsal segment barely attaining the middle of the tooth 
when tibia is flexed. Mesothoracic femur one-fourth longer than the tibia, 
which is two and one-half times as long as the tarsus without the claws. 
Metathoracic femiu' and tibia of equal length, the tibia about three' and one- 
half times the length of tarsus without the claws, which are one-third the 
length of the tarsus. 

Notes. The above species is described from a series of 20 speci- 
mens, 16 of which were collected by Mrs. Grace Wiley in Colorado 



Hl\ni;i:kfori): Xepid.k in Amkkica. 433 

county, Texas. Three were taken in New Orleans, La., and belong 
to Dr. Carl Drake; and one from tlie state of Coloma, Mexico, is in 
the United States National Museum. The species is named in honor 
of Dr. Carl Drake, who believed the species to be new as long ago 
as 1916. but who nc\-er found time to study the matter thoroughly. 

Cvricta drakeij while measuring as long as C. houmrdii, is a much 
more slender species, and appears smaller. Indeed, comparing the 
males of the two species, there is a considerable difference in size 
and shape. The anterior lateral lobes of the pronotum are much 
more prominent in C. houmrdii, the sides of prothorax, therefore, 
more curved. The median longitudinal fossa in C. howardii is con- 
fined to anterior part of the median elevation. The tylus is longer 
in C. drakei, the antennse are differently formed (see figures 7 and 8 
on plate XLVII), and the mietasternal plate is smaller. This new 
species differs from C. voire tni Montd. from Mexico by its smaller 
size (C volxemi is 241^ mm. long without appendages), by the tooth 
on the front femur being conspicuously nearer base than apex, while 
Doctor ]\Iontandon's figure of his species shows the tooth in the 
middle, or nearer the apex. It further differs from C. volxemi by 
having the head longitudinally carinate and the scutellum tricari- 
nate. The thorax is relatively shorter than in C. volxemi, which 
has a thorax twice as long as wide. The front coxae of C. drakei are 
much shorter. 

It differs from C. scorpio Stal, which, according to Montandon, 
lacks the carina on head and scutellum and has the sides of the 
thorax much as in C. howardii Montd. The longitudinal grooves of 
the thorax of C. scorpio Stal are less emphasized than in C. howardii, 
which in turn has them less emphazised than C. drakei. All of these 
points separate C. drakei from C. scorpio Stal, and are based upon 
Doctor Montandon's remarks upon the latter species. Champion, 
in his "Biologia Central! Americana," places Nepoidea montandoni 
Martin as a synonym of C. scorpio Stal. Joanny Martin, under the 
title "Descriptions d'Especes Nouvelies de Nepidse (Hem.)," in Bul- 
letin de la Societe Entomologique de France, 1898, pages 66-68, de- 
scribes on page 68 his A^. montandoni, and figures the head and 
thorax on the previous page. C. drakei is unlike this species. Stal's 
description of C. scorpio is wholly inadequate. 

Genus Ranatra Fabricius 1790. 
This genus is characterized by its very elongate, slender form. 
It is the dominant genus of the family Nepidse in North America. 
Doctor Horvath, in his paper, "Les Relations entre les Faunes 



434 The University Science Bulletin. 

Hemipterologiques de I'Europe et de I'Amerique du Nord" (Pro- 
ceedings of the Seventh International Zoological Congress, 1907), 
lists the genus Ranatra as belonging to the oriental fauna. Since 
that time Doctor Montandon has added several names to our Ameri- 
can lists. 

The species in the genus are superficially so similar in color and 
general characteristics that there has been much confusion in 
naming them. The descriptions have been made in several cases 
from a single specimen, without, therefore, a consideration of varia- 
tions within the species. They have dealt with comparisons that 
can be appreciated only after long familiarity with the insects, and 
on this account many errors of determination have arisen. The 
most unfortunate circumstances attending the study of North 
American Ranatra has been the failure to recognize the identity of 
Ranatra jusca P. B. 1805 and Ranatra nigra H. S. 1853, the de- 
scriptions of which are inadequate, and therefore, since the types are 
not available, can be interpreted authoritatively only through the 
examination of abundant and representative material from the 
entire country. 

After studying long series of specimens representing a wide dis- 
tribution, the writer has been compelled by the evidence before him 
to place a new interpretation upon our North American Ranatra. 
This lie has been reluctant to do, because it involves the renaming 
of our commonest two species. Stability, however, can never be 
attained in this group by postponing or ignoring the evidence that 
eventually must prevail. 

Ranatra jusca P. B. was described and figured in color, natural 
size, by Palisot de Beauvois, in his 'Tnsectes recueillis en Afrique et 
in Amerique, dans les Royaumes d'6ware et de Benin,a Saint Domin- 
que et dans les Etats Unis, pendant les annees 1786-1797." 

The figure is not amplified nor enlarged as suggested by Doctor 
Montandon (Bulletin Soc. Sci. Bucharest, vol. XIX, 1910), who was 
in error in endeavoring to make R. kirkaldyi Bueno, our smallest 
North American Ranatra, fit a drawing that has the dimensions of 
our largest species. He was mistaken in believing R. kirkaldyi 
Bueno a synonym of R. jusca P. B., for there are two convincing 
reasons for believing R. jusca P. B. is represented natural size. 

First. Palisot de Beauvois in his "Discours Preliminaire," page 
XV, says, "J'ai adopte, pour la grandeKur des figures, un plan uni- 
forme, et qui m'a semble'plus commode, c'est-a-dire,celle d'un pouce 
pour tous les Insectes plu^ petits que cette mesure adoptee, en plac- 



Hungerford: Nepid.k in America. 435 

ant a cote une ligne perpendiculaire- de la grandeur reerdc Tanimal. 
Ceux (lui excedent un ponce seront figures de Icur grandeur na- 
turelle." 

Second. The plate which bears the figure of Ranatra fusca has 
upon it three figures natural size (one of which is Ranatra fusca)' 
and two figures enlarged. These two have the true length indicated 
by a line, and one of these, Naucoris jemorata, of w^iicli the writer 
has specimens, fits the line which was drawn to indicate the true 
length. There are on the plate three figures of Belostoma which 
Palisot calls "Nepa," male and female, of "Nepa suhspinosa," nat- 
ural size, which are not accompanied by a "line of true length," and 
Neya minor, accompanied by a line. The drawing of Neya minor is 
as large as those of Nepa subspinosa, and yet the author distinctly 
says that Nepa minor is smaller. The line indicates its true length, 
and the inevitable conclusion is that the figure of Ranatra fusca is 
natural size. Indeed, a comparison of drawings with the specimens 
throughout the book shows the author consistently followed his 
stated policy of drawing large insects natural size and indicating 
enlargements by lines, whenever made. 

Palisot's insects were taken between the years 1786 and 1797. 
Those from the United States, then, were taken from somewhere in 
the eastern United States, for the territories of the United States at 
that time were bounded on the north by Canada, on the west by the 
Alississippi river, and on the south by the north line of Florida. 
Ranatra fusca, therefore was collected within these limits. 

Palisot, in his "Discours Pr^liminaire," alludes to an account of his 
travels as being in press. I have not located this, but have secured 
two brief biographical sketches of him; one from "La Grande Ency- 
clopedie," vol. XXV, and the other more complete, from P. Larousse, 
"Dictionnaire du XIX Seicle 12," page 66. Neither of these mention 
a book of his travels. From these accounts we learn that he came to 
Philadelphia first in 1791 to secure assistance against a Negro up- 
rising in San Domingo, where he was taking part in govermental 
affairs. He seems to have been in the United States for some time, 
for upon his return to San Domingo he narrowly escaped death at 
the hands of the blacks (17931. He escaped to Philadelphia with- 
out funds and supported himself there by giving lessons in music 
and language. Finally he secured help from France to make a 
scientific voyage in North America for new collections, and returned 
to his own country in 1798. He died in 1820. These brief facts 
indicate that most of his days in the United States were spent about 
Philadelphia, and it seems to me probable that many of his insects 



436 The University Science Bulletin. 

were collected in that region. The possibility of his R. fusca not 
being known to us in nature seems to me quite remote. 

According to the figure, Ranatra fusca of Palisot de Beauvois is 
a large, robust insect with broad anterior femora and short legs and 
respiratory tube. The cjuestion is, what insect of the range indi- 
cated has these characters? After careful consideration of the 
problem I must conclude that the only one with the proper size, 
robustness, broad femora and short legs is Ranatra americana 
Montd. It has the variable respiratory tube, usually longer than 
Palisot's figure, but not uncommonly as short, and in every case 
shorter than the body. The failure of the artist to indicate any 
apical tooth on the anterior femur is readily understood, since we 
know that this character is often obscured by a marginal fringe of 
pile so completely as to escape any but the closest scrutiny. It is 
indeed a somewhat variable structure; in some examples much re- 
duced, and in others plainly visible. The writer has in his collection 
specimens which fit Palisot's figure almost exactly. 

The second species with which we are concerned is Ranatra nigra 
H. S. This species was described in 1853 from America by Herrich 
Schaffer in his "Die Wanzenartigen Insecten." It was described as 
being from 2 to 21/8 inches long from beak to tip of respiratory 
tubes, with the respiratory tubes not much over half as long as the 
body; indeed, he says they were three-fourths of an inch! Now the 
only species we have which in a series of specimens has a respiratory 
tube averaging three-fourths of an inch long is R. protensa Montd. 
Doctor Montandon described his R. protensa from a single large 
female, which, because of its very yellowish color and shorter limbs, 
did not suggest R. nigra to his mind. Indeed, considered alone, it is 
not at all to be expected that it would. R. nigra was described as 
having the structure of thorax and relative length of limbs as in i?. 
elongata. Now, R. elongata has very elongate hind femora, the tips 
surpassing the last abdominal suture by a considerable distance. 
The length of limb in R. protensa Montd. is, on the whole, not as 
great as in R. elongata, but is strikingly longer than in R. linearis 
L., with which he compared it in size, and this suggested R. elongata. 
Indeed in many specimens of R. protensa Montd. the hind femora 
surpass the last abdominal suture. The front legs of R. protensa 
are long and very slender and the thorax is more like that of R. 
elongata than of R. linearis. Doctor Montandon said that he had 
never seen any specimens from North America with legs as in 
elongata. We have three species with limbs relatively as long. 



Huxcerford: Nepid.i-: ix America. 437 

It seems scarcely necessary to mention that the color implied by 
the name 7iigra has nothing to do with the case, for black coloration 
occurs in all the species, due to one of two causes — either dark in- 
crustations or deposits upon the integument, or dark discoloration 
due to failure in drying out the specimens. Either of these causes 
might account for a uniform dark or black color in a given series 
of insects. 

There are other facts relative to the identity of these species 
which will be presented under notes after the various descriptions, 
to which the I'eader is referred for further evidence. Aside from the 
points mentioned, the probability of our two commonest and wide- 
spread species being first described is of itself very great. 

The list of the species of Ranatra now known from America north 
of Mexico is as follows: 

Ranatra fusca P. B. 1805 = Ranatra americana Montd. 1910. 

Ranatra nigra H. S. 185^ ^ Ranatra protensa Montd. 1910. 

Ranatra quadridentata Stal 1861. 

Ranatra kirkaldyi Bueno 1905 and its variety hofjmanni, n. var. 

Ranatra brevicollis Montd. 1910. 

Ranatra buenoi, sp. new = Ranatra fusca Bueno and Mondt. in 
part. 

Ranatra drakei, sp. new. 

Ranatra australis, sp. new. 

It will be noted that R. quadridentata Stal, completely submerged 
in Van Duzee's catalogue under R. americana Montd., is restored. 
Dr. F. H. Snow, in Trans. Kan. Acad. Sci., vol. XX, pt. 1, p. 153, 
1906, w^as not writing about the species which Doctor Montandon 
later described as R. americana, but another which we believe to be 
R. quadridentata Stal, and should not have been synonymized by 
Van Duzee. It will also be seen that R. annulipes Stal is omitted. 
This R. annulipes Stal is a very distinct species and not to be mis- 
taken for any the writer knows from our range. Doctor Montandon 
has established, through the examination of types and type ma- 
terial, that R. fabricii Guer. is the same as R. annulipes Stal. Fig- 
ures of this species are given in this paper, because it has been cited 
as coming from our range. 

The variety, edentata Montd., of R. americana Montd., is as- 
sumed to be an americana with attenuated apical tooth on front 
femur. Whether it should be recognized as a variety is questionable. 

The entire question of the correct names for the Ranatra was, as 
the writer has stated, opened by the necessity of naming four species 



438 The University Science Bulletin. 

from Kansas. The problem was undertaken with no preconceived 
notions in the matter, except, indeed, a leaning toward accepting 
Doctor Montandon's studies, since he is a great scholar in the 
Cryptocerata and was followed by Van Duzee in his catalogue. The 
results herewith presented may be received with some irritation, 
because of the revolution necessary in nomenclature, but it seems 
best for us to get as near the truth as the evidence permits. 

In an endeavor to fix the limits of the species, the writer has 
studied the characters used by the best systematists in the group 
and in addition, antennal and genital structures. The relative length 
of body parts, the comparative size of the eyes, the relative length 
of the limbs and of the caudal filaments, the shape of the sternum 
and the form of the front femora have been in general use in the at- 
tempts to define nature's species in this genus. Species are expected 
to vary within limits, and breeding experiments with insects show 
them to vary, not only in single characters, but in linked characters, 
and in order to steer a fair course between ''lumping" and ''split- 
ting" species, it is much better to use a combination of three or more 
characters than to depend too much upon any single one. The use, 
therefore, of the characters of the antennae and genitalia is a ma- 
terial aid in the classification of this difficult group. 

NOTES ON TERMS USED IN KEYS AND DESCRIPTIONS. 

The apical tooth of the front femur is on the inside edge of the 
front femur near the attachment of the tibia (see fig. 2-A, pi. 
XLIV). The antennae are hidden in pockets beneath the eyes (see 
fig. 9^ pi. XLV). The prothorax of Ranatra is divided into two 
parts, anterior and posterior, by transverse lateral grooves (see fig. 
2-G, pi. XLIV). The tylus and jugae are figured on plate XLV, 
figure 12. The metaxyphus extends back between the hind coxae 
as a part of the metasternum. The last abdominal segment is desig- 
nated as the "genital segment," and its ventral plate in both sexes as 
the "operculum." 

In this paper I have not distinguished between the genital seg- 
ment and the last abdominal segment, as is often done with some 
other Heteroptera, where the last abdominal segment is considered, 
the segment just in front of the genital segment. I refer to this as 
the penultimate abdominal segment (see figs. 10 and 11, pi. XLV). 

The male genital capsule of which I speak is, I take it, homolo- 
gous with what G. C. Crampton in the Bulletin of the Brooklyn 
Entomological Society, volume XVII, pages 45-55, calls the hypan- 
drium," and the claspers correspond to his "styli," or "gonostyles." 



Hungerford: Nepid.e in America. 439 

TECHNIQUE USED. 

The antennee arc often covered with debris that should be re- 
moved. If the antennae are still difficult to see, the specimen may be 
relaxed and antennae moved into view with dissecting needle. The 
genital capsule of the male lies in the last abdominal segment, en- 
tirely hidden. To examine this, relax the specimen in a moist cham- 
ber, then holding the specimen on its right side in the left hand, 
lower the operculum with a dissecting needle and draw out the 
capsule. A needle with just the very tip turned at right angle is 
best to use. It can be inserted alongside the capsule, turned so 
that the hook is against the capsule, and slowly withdrawn, bringing 
with it the desired organ. If the respiratory filaments start to come 
also, loosen the capsule from them by inserting the needle between 
capsule and filaments on each side. The capsule may be removed 
entirely and mounted upon a card below the insect, or simply drawn 
into view. The insects are not in the least damaged, and the male 
capsules may be taken out and operculum pressed back into place 
by a careful person without it being detected by the closest exam- 
ination of the exterior. 

KEY TO THE SPECIES. 

A. Antennae simple, distal end of the penultimate segment without lateral 
prolongation. Front femora broad and stout and not narrowed near 
middle. 

B. Front femora without apical tooth or marked sinuosity. 

R. kirkaldyi Bueno. 
BB. Front femora with apical tooth or a marked sinuosity. 

R. kirkaldyi Bueno. 
var. hoffmanni, new. 

AA. Antennae with distal end of penultimate segment with a lateral pro- 
longation. Front femora somewhat narrowed in their middle section. 

B. The lateral prolongation of the penultimate segment of- antennae 
not greater than one-half of the length of the ultimate segment. 
Front femora very slender and without apical tooth. 

C. Prosternum with a single wide, deep longitudinal trough. Eyes 
veiy prominent. . R. buenoi, sp. new. 

{=R. jusca Bueno.) 
CC. Prosternum without the deep trough, but possessing two 
longitudinal depressed lines characteristic of most species of 
Ranatra. R. nigra H. S. 

{=R. protensa Montd.) 
BB. The lateral prolongation of the penultimate segment of antennae 
greater than one-half the length of the ultimate segment. 
C. Sides of body (connexivum) embracing the operculum of the 
genital segment at its extremity (see fig. 11, pi. XLV), 

R. annulipes Stal. 
{=:R. fabricii Gu^r.) 



440 The University Science Bulletin. 

CC. Sides of body not embracing the genital operculum at its ex- 
tremity (see fig. 10, pi. XLV). 
D. Pronotum broad and short, anterior enlargement sub- 
equal in width to entire head. Anterior femur broad and 
stout without apical tooth. Median tooth half way be- 
tween trochanter and tibial joint (measured on inner 
edge). Metaxyphus very long, extending beyond middle 
of hind coxae, nearly attaining abdomen. 

R. brcvicoUis Montd. 
DD. Pronotum more slender. The median tooth of anterior 
femur nearer distal than proximal end. Metaxyphus 
not often extending bej'ond the middle of hind coxae. 

E. Front femur broad without apical tooth. 

F. Jugae of head more prominent than tylus. 

R. mistralis, sp. new. 
FF. Tylus fully as prominent as jugae. 

R. jusca P. B. 
(var. edentula Montd.) 
EE. Front femur with apical tooth. 

F. Anterior portion of prothorax fully twice as 
long as thickened posterior portion. Eyes 
large, plainly greater than interocular space. 
Jugae of head prominent. Front femur long 
and slender. Hind femur surpassing middle of 
penultimate abdominal segment. Caudal fila- 
ments as long as body. R. drakei, sp. new. 
FF. Anterior portion of prothorax shorter than 
above. Eyes not much, if any, greater than 
interocular space. Jugae not so prominent. 
Front femur fairly stout. Middle and hind 
femora short, not attaining middle of penulti- 
mate segment. 

G. Eyes prominent, plainly transverse, fully 
as large as interocular space. Anterior 
part of pronotimi somewhat flattened 
dorsoventrallv. Hind femora short. 

R. jusca P. B. 
(=:R. americana Montd.) 
GG. Eyes not prominent, neither transverse 
nor as large as interocular space. An- 
terior part of pronotum more cylindrical. 
R. quadridentata Stal. 

. Ranatra kirkaldyi Bueno. 

Bueno, J. R. de la Torre, Can. Ent. XXXVII, p. 187, 1905. 

(= -R. fusca Montd., Bui. Soc. Sci. Bucharest, XIX, p. 3; 1910.) 

Original description: 

Abdominis dorsum orange brown; eyes small, not very prominent; pro- 
thorax much constricted at the middle, bisulcate beneath; wings smoky; 
anterior femora broad with a prominent tooth near the middle, otherwise 
smooth; posterior tarsi extending beyond the middle of the air tube; air 
tube shorter than the length of the abdomen; legs banded. Length from tip 
of abdomen to tip of rostrum, male, 23 mm. to 26.4 mm.; female, 27 to 31 mm. 



Hungerford: Nepid.e in America. 441 

Notes. Tliis is the species that Doctor Montandon has con- 
sidered a synonym of I\\ fusca P. B.* Its much smaller size (7?. 
fusca P. B. measures 37 mm.) is not the only reason for believing 
otherwise. Tlie anterior femora of R. kirkaldyi Bueno are not con- 
stricted near the middle tootli as they are in R. fusca P. B. The 
front legs and respiratory tube are both shorter relatively than in 
Palisot's figure. This is the smallest known species in North Amer- 
ica. It is a characteristic species, quite unrelated and distinct from 
the others. In the first place, the antennae are much reduced. They 
are straight, lacking any projections from the penultimate segment, 
and often lack anything more than a constriction to distinguish the 
terminal segment. The ultimate segment is sometimes fused with 
the penultimate (see figs. 9 and 11, pi. XLVII), but all stages of 
separation can be found. The male genital claspers are also very 
distinct, as shown by figure 5 on plate XLVI. 

Specimens of this species taken by Beamer and Hungerford in 
Cherokee county, Kansas, were identified by Mr. J. R. de la Torre 
Bueno. They agree with the original description in lacking any 
apical tooth or sinuosity near the apex of the femora. They measure 
from tip of beak to base of filaments, from 24 to 30 mm., wdth the 
filaments from 14 to 17 mm. long. I have seen the true R. kirkaldyi 
Bueno from New York. 

In addition to the above Kansas series, I have before me the 
following series of 127 specimens as follows: 

8 Rockbridge, Ohio, 9-30-16, C. J. Drake. 

1 Rocky Mount, N. C, 10-19-16, R. W. Leiby. 

3 St. Paul, Minn., Phalen Lake, 6-19-21, H. B. Hungerford. 

1 St. Paul, Minn., Lake Johanna, 6-26-21, H. B. Hungerford. 

4 Ramsey county, Minnesota, 1-9 22, W. E. Hoffman. 

7 St. Paul, Minn., Lake Johanna, 10-13-22, W. E. Hoffman. 

9 St. Paul, Minn., Lake Johanna, 10-14-22, W. E. Hoffman. 
32 St. Paul, Minn., Lake Johanna, W. E. Hoffman. 

3 Lincoln, Neb., 1 male, 2 females, W. E. Hoft'man. 

39 St. Paul, Minn., Water Supply Canal, 10-18-21, W. E. Hoffman. 
16 St. Paul, Minn., Lake Johanna, 9-24-21, W. E. Hoffman. 

4 St. Paul, Minn., Lake Johanna, 9-17-21, W. E. Hoffman. 

I have also seen a series of 170 taken by Doctor Drake in Ohio. 

These Minnesota specimens have certain characteristics which 
set them clearly apart from the Kansas series, yet there can be no 
mistake in considering them as belonging to the same species. Since 
I have had a fairly good series of these insects, which are smaller 

*See plate XLIX, figures 1 and 2. 



442 The University Science Bulletin. 

than the Kansas forms, more robust, with front femur relatively- 
thicker and possessing a more or less well-marked sinuosity near the 
inner apex, with antennae which, on the whole, are blunter at tip, 
and the subapical tooth of the claspers of male slightly more slen- 
der, it seems wise to consider them a variety, for which I propose 
the name Ranatra kirkaldyi var. hoffmanni. It will be noted that 
Mr. W. E. Hoffman has provided me with the longest series of this 
variety. In a series as large as this, it would seem that if the variety 
were not fixed there should be found forms like the true kirkaldyi, 
which is described as having "anterior femora broad with a promi- 
nent tooth near the middle, otherwise smooth." 

Ranatra buenoi, sp. new. 

= R. fusca Bueno, Can. Ent. XXXVII, p. 188, 1905. 

Bueno and Brimley, Ent. News, XVIII, p. 438, 1907. 

Bueno, Ent. News, XXXII, p. 273, 1921. 

Montandon ; determination in U. S. National Museum. 

Size. Length from tip of beak to tip of abdomen, 32 mm. to 38 mm.; caudal 
filaments, 22 mm. to 27 ami. long.. 

Color. From light to very dark fuscous; top of abdomen orange and black; 
middle and hind legs of lighter forms banded. 

Shape. Long and slender, prothorax long and slender; eyes very prominent 
and large; limbs all very slender and very long; the under side of prothorax 
with a single broad and deep sulcus, which distinguishes this species from any 
of our other forms (see fig. 6, pi. XLVIII). 

Structural peculiarities. The eyes very prominent, transverse diameter 
greater than the interocular space; tjdus longer than jugae and as prominent 
as these parts, which are of medium development; antennae with the lateral 
prolongation of penultimate segment not more than half the length of ulti- 
mate; prothorax slender, the anterior portion measured on the median dorsal 
line, twice the length of the posterior swollen part (the well-marked long trans- 
verse lateral grooves used as di\ading line) ; scutellum somewhat elevated and 
slender and the length of the abdomen is to pronotum as 2^ is to 1; respira- 
tory filaments surpassing front margin of scutellum when brought forward; 
claspers of male genital bulb very distinct; subapical tooth greatly reduced 
(see fig. 1, pi. XLVI) ; all the limbs strikingly long and slender, especially the 
anterior femora (see fig. 1, pi. XLVIII) ; no apical tooth on anterior femur, the 
other tooth much nearer the apex than the base, the basal part being at least 
IV2 times as long as the part lying before the tooth; the coxa two-thirds as 
long as the femur. 

The middle and hind legs long, the distal ends of hind femora surpassing 
the last abdominal suture and often nearly attaining caudal end of genital 
segment, while the middle femora frequently attain or surpass the caudal 
margin of penultimate abdominal segment; the distal ends of middle and 
hind legs almost attaining tip of respiratory filaments; the relative lengths 
of femora to tibiae are about as 16 is to 19 for the middle leg and as 16 is to 
22 for hind leg. The tarsi are small relatively, a little less than one-sixth as 
long as their tibiae. 



Hungerford: Nepid.k in America. 443 

Notes. Described from the following series: 

Male holotype, Colorado county, Texas, June 24, 1922; Mrs. Grace Wiley, 

collector. 
Female allotype, Colorado county, Texas, June 24, 1922; Mrs. Grace Wiley, 

collector. 
Paratypes as follows: 

2 9 $ Raleigh, N. C, July 10, 1902; F. Sherman, jr., collector. 

$ Mound, La., November 6, 1918. 
9 Aberdeen, Miss.; Dr. Carl Drake. 

3 9 9 and IS Leiand, Miss., September 16, 1921; C. J. Drake. 

9 Creve Coeur Lake, Mo., May 15, 1911; J. F. Abbott. 
$ Dime Box, Tex., July 20, 1911; C. T. Atkinson, collector. 
17 ,J 5 and 22 9 9 Gainesville, Fla., June 19, 1918; Carl Drake. 

Holotype and allotype in University of Kansas museum; para- 
types in collection of Dr. Carl Drake, J. R. de la Torre Bueno, and 
the author. 

This species has been named for Mr. J. R. de la Torre Bueno, 
who pointed out its structural characters in 1905.* It has also 
been named by some workers R. fusca P. B., and by at least one 
R. nigra H. S. It is clearly impossible for it to be either. The error 
of considering it R. fusca P. B. has been due to the inadequate de- 
scription of R. fusca P. B. and to the fact that the original text with 
illustration has not been accessible to many, if any, American stu- 
dents of this group. By taking Palisot de Beauvois' figure of his 
R. fusca and comparing it with the species above described, it will 
be seen at once that the species are not the same. The long, slender 
limbs, the great eyes, the long thorax, the relative position of the. 
tooth on the front femur, and the relation in length between femora 
and tibiae of the legs, preclude the possibility that Palisot's artist 
had R. buenoi, sp. new, before him. The front legs of R. fusca P. B. 
are stout, not exceedingly slender. The median tooth of front femur 
is nearer the middle than in R. buenoi, sp. new. The anterior part 
of prothorax is less than twice the swollen part. The middle femora 
and tibise are nearly the same length as they are in R. americana 
Montd., for instance, and not considerably different in length as in 
R. buenoi. The hind femora are considerably shorter relative to the 
abdomen in R. fusca P. B. The tarsus of R. fusca P. B. is larger, 
being one-fifth as long as its tibia and not one-sixth as in R. buenoi 
sp. new. In other words, its proportions fit another American spe- 
cies, but not R. buenoi. 

This species is not R. nigra H. S., for R. nigra H. S. has uni- 
formly short respiratory tube of three-fourths inch and measures 

*See Can. Ento., vol. XXXVII, p. 188; 1905. 



444 The University Science Bulletin. 

over all from tip of beak to end of filaments 2 to 2% inches, while 
this species has a tube averaging a full inch and varies in size from 
2^8 to 21/2 inches. The shape of the eyes as shown in Herrich- 
Schaffer's figure, is very different, and the size which he says is 
"not larger than R. linearis," would at once eliminate it, for in 
R. buenai they are very large, but not so large as in R. elongata, 
specimens of which I have seen. 

For illustration of this species, see figures 1 and 2, plate XLVI; 
fig. 12, plate XLVII; figure 1, plate XLVIII; and figure 5, plate 
XLIX. This species, as Bueno has written, is southern in distribu- 
tion. Besides the localities above cited, I have seen it from Way- 
cross, Ga., taken by J. C. Bradley, U. S. N. M. 

Ranatra nigra Herrich-Schaffer. 

Henich-Schaffer, G. A. W., Wanzenartigen Insecten, IX, p. 32 ; 1853. 
(^ Runatra proie»sa Montd. and all subsequent writers.) 

Original description : 

R. nigra m. tab. 290, fig. L, 2 = 2%". Nigra, tubis respiratoribus longi- 
tiidinem dimidiam corporis parum superantibus. Grosse fast von R. linearis, 
schwarzer, die Aiigen night grosser aber seitlich mehr vortretend, der Scheitel 
daher breiter. Ban des Thorax imd Liingenverhaltnisse der Beine wie bei 

R. elongata. Aus Anierika. 

In addition to the above, and in the discussion relative to the 
genus Ranatra and the species R. elongata, R. filiformis, R. linearis, 
macrophthalma and R. nigra, he gives notes of comparison which 
throw further light upon his species. He says R. elongata has the 
longest respiratory tube (134 inches) and R. nigra has the shortest 
{% inch). The color is darkest in R. nigra, "die Augen sind am 
grossten bei macrophthalma, am kleinsten bei linearis, am rundes- 
ten bei letzerer, am meisteiijaie Quere gezogen bei nigra." His state- 
ment of length, 2 to 2^8 inches from tip of beak to tip of tube with 
"Athmungsrohren nicht viel iiber halb so lang als der Korper," in- 
dicate that he had a series of the insects. 

Notes. The only species of our country which agrees in size and 
uniformly short length of respiratory tube with the description of 
R. nigra is the insect we know under the name R. protensa Montd. 
Doctor Montandon had before him a single large, fairly short- 
limbed female, which is his type deposited in the United States Na- 
tional Museum. This $ was collected by Wm. H. Ashmead on 
Long Island. The respiratory tube is shorter than the abdomen. 
The legs extended would surely reach the tip of the respiratorv tube. 

Ranatra nigra is quite common over our country everywhere I 
have collected. It is most unfortunate that it was described from 



Hungerford: Nepid.e in America. 445 

blackened specimens, for indeed it is our lightest-colored species, 
being of a yellowish color with a trace of green throughout. This 
species has very prominent mesocoxae and metacoxse, which are 
angular, having a slight tubercle on the inner side. The breast, or 
prothorax in front of the mesocoxal elevations, is constricted and 
small, making the coxal elevations very pronounced. 

The United States National Museum possesses, besides the type, 
specimens from Virginia, West Virginia, Pennsylvania, Maryland, 
Arkansas and Florida. 

I have before me a series of seventy-nine specimens from Doni- 
phan county, Kansas, taken by Robert Guntert and W. J. Brown; 
ten from Douglas county, Kansas, and forty specimens taken by 
Mr. W. E. Hoffmann and myself from the following places in 
Minnesota: Lake Johanna, Phalen lake and Minnehaha creek, all 
near the twin cities. We have also taken it in other parts of the 
state. Careful study of ample material, well distributed, shows 
that by far the commonest size is, total length, two inches, with re- 
spiratory tube three-fourths inch! The length of hind femur rel- 
ative to abdomen varies, sometimes attaining the front margin of 
the genital segment and sometimes falling somewhat short of this. 
The eyes are transverse and somewhat protuberant. The jugse fit 
closely against the tylus. The lateral prolongation of the penulti- 
mate segment of the antennae is less than half the ultimate. The 
front femora are very slender and coxae long. On page 156 of the 
University of Kansas Science Bulletin, vol. XI, where I give an 
English translation of Doctor Montandon's description in French 
of R. protensa, I should have written, "Anterior femora quite slen- 
der, but scarcely a fifth longer than their coxae," instead of "as long 
as" — a careless error. 

Ranatra annuli'pes Stal. 

Sfal, Of. Vet. Akad. Forh., XI, p. 241, 1854. 

Original description: 

R. anmdipea : Flavotestacea; hemelytris fuscescente testaceis; pedibus obso- 
lete fuscoannulatis; spiraculis nigrofuscis. Long. 30, lat. 3% millim. — Brasilia. 

In 1861, under the title "Genera Nepidarum synoptice desposita," 
in his "Nova methodus familias quasdam Hemipterorum dispon- 
endi," "Ofversigt af Kongl. Vetenshaps akademiens forhandlingar, 
Arg. 18, 1861, No. 4, he gives a more satisfactory description: 

Ranatra annulipes. Stal. Nepidse, 1861, p. 9. 

R. annulipes. Pallide testaceo-grisea, pedibus obsolete fusco variis; abdo- 
mine dorso sanguineo-fusco, lateribus griseo-flavescentibus, stigmatibus nigris, 



446 The University Science Bulletin. 

dorso pellucentibus ; fronte convexiuscula ; alis levissime infuscatis; prosterno 
bisulcato; femoribus anticis pone medium intus unidentatis, extus ibidem 
at prope apicem inermibus, intermediis posticis subbrevioribus, his ad medium 
segmenti ultimi abdominis porrigendis; aidothecae appendicibus corpori sequi- 
longis. Long. 30 millim. Brasilea (Mus. Hohn.). 

Ranaira annulipes Stal. Of. Vet. Ak. Forh., 1854, p. 241-1. Prsecedentibus 
duabus affinis et cum iisdem divisionem forma metasterni distinctissimam, 
Americam habitantem formans. Secundum formam metasterni in divisiones 
quattuor distinctas, determinationem specierum facilitantes dividi potest 
Ranatra genus." 

Doctor Montandon (Bui. Soc. Sci. Bucharest, XVIII, 1910) , who 
has examined Guerin's type of R. fabricii (1857) pronounces it to 
be the same as R. annulipes Stal, 1854. Sagra's Historia fisica, po- 
litica y natural de la Isla de Cuba, volume VII (1857) is not avail- 
able to many of our workers; therefore, Guerin-Meneville's descrip- 
tion of R. fabricii follows: 

Ranatra fusca tubo respiratorio corpora paulo longiore ; pedibus anteriori- 
bus nigro-fuscis ; corpoie infra obscure feiTUgineo. Larg. 35, Anch. 3 millim. 

Notes. This species is distinguished from all our species by hav- 
ing the connexivum of the genital segment enlarged and extended 
ventrally at the caudal end embracing the distal portion of the geni- 
tal operculum (see fig. 11, pi. XLV). I have never seen specimens 
of this species from the United States and Prof. H. G. Barber in- 
forms me that his record from Texas was on a mistaken identifica- 
tion. Mr. W. L. McAfee says the species appears to be common 
in Cuba. 

Ranatra fusca Palisot Beauvois. 

Palisot de Beauvois, A. M. F. J., Ins. Rec. Afr. Am., p. 235; 1805. 
(= R. Americana Montd., 1910, and subsequent writers.) 

Original description: 

RanaUre brune. Brune-verdatre ; soies un peu plus courtes que le corps; 
ailes brunes-roligeatres. (Fig. 1.) 

Ranatra fusca. V«ridi-f usca ; setis corpore brevioribus; alis fusco-rubellis. 
(Fig. 1.) Etats-Unis d'Amerique. 

Obs. Cette espe^p differe de celle d'Europe par sa couleur plus pale, par les 
soies qui terminent I'abdomen, plus courtes que le corps, par les ailes et le 
dessus de I'abdomen d'un brun rougeJitre. 

Notes. A photographic reproduction of Palisot's figure (natural 
size) is given on plate XLIX, figure 1. On a previous page I have 
stated the evidence to show that Palisot's illustration is natural size. 
Size alone shows that Palisot's species is not the same as R. kir- 
kaldyi Bueno. It also eliminated R. nigra H. S. {= R. protensa 
Montd.). The short legs and stout front femora eliminate R. fusca 



Hungerford: Nepid.k in America. 447 

Bueno. Even after making due allowances for "artist's license," 
it is not conceivable tliat the careful artist illustrating Palisot's spe- 
cies could have been looking upon R. fusca Bueno. The eyes of R. 
fiisca Bueno are strikingly large, the anterior femora very slender, 
and the middle and hind tibia? much larger in proportion to their 
femora than in Ranatra fusca P. B. 

Doctor Montandon (Bui. Soc. Sci. Bucharest, XIX, 1910), who 
also has examined Palisot's figure, says in effect regarding it: ". . . 
When one considers only the essential details for the characteristics 
of a form, such as the proportional length of the legs and append- 
ages, one recognizes without difficulty its short and very robust an- 
terior femora, its very little developed posterior legs, and its ap- 
pendages shorter than the abdomen . . ." 

He was led into error by assuming the figure to be a great en- 
largement of Bueno's R. kirkaldyi, which cannot be true according 
to the facts elsewhere presented in this paper. Doctor Montandon 
says that his R. americana is readily distinguished by the shape of 
the prothorax, and indeed he is correct. The posterior enlarged part 
is plump and then narrows to a slender neck, then widens again in 
front. Any student with much experience with American Ranatra 
will recognize this characteristic at once (see Palisot's figure repro- 
duced on plate XLIX, figure 1). Compare it with the photographs 
of our other species. Palisot's insect was broad and large; our only 
species comparable to it is R. americana, which is as large, some- 
times a trifle larger. The anterior portion of prothorax is about 
one and one-half times the swollen posterior portion; so it is in R. 
americana Montd. (in R. fusca Bueno [= R. buenoi, sp. new] the 
anterior part is about twice the posterior part). The legs are short; 
so with R. americana Montd. The large ratio of length of tarsus to 
the tibia, 1 to 5+ (in R. fusca Bueno, 1 to 6+), shows similarity; 
also the hind tibia is a little longer than the femur, but not as much 
as in R. fusca Bueno. The broad anterior femora agree with R. 
americana, but the apical tooth is not shown. This, however, is a 
character which all students of this group realize could be overlooked 
readily. Sometimes the space in front of the tooth is filled with 
debris, obscuring the tooth; sometimes the tooth itself is much re- 
duced. Doctor Montandon has a specimen from Philadelphia and 
another from Texas which lack the tooth and which he designates 
under the name R. americana var. edentula. In many of the speci- 
mens the tooth, therefore, is not marked and must have been over- 
looked by Palisot's artist. The caudal filaments are comparatively 



448 The University Science Bulletin. 

short in the figure; so they are in many specimens of R. americana. 
I possess examples with even shorter filaments. On the whole, I 
should say the average is longer, but not as long relatively as in i?. 
buenoi, sp. new (= R. fusca Bueno), or in R. drakei, sp. new, or in 
R. australis, sp. new. It is most robust and, on the whole, our larg- 
est species. I have collected it in New York, Minnesota and Kansas 
in numbers, and have seen specimens from various other states. 

Ranatra brevicollis Montandon. 

Montandon, A. L., Bui. Soc. Sci. Bucharest, XVIII, p. 184; 1910. 

Original description: 

Ranatra brcuicoliis nov. sp. C'est bien a regret que je decris cette nouvelle 
forme sur un exemplaire malheureusement unique, assez peu dissemblable, au 
premier aspect de R. quadridentata Stal, mais ses caracteres specifiques ne 
premettent pas de la confondre avec les autres especes fusca ou quadridentata 
dont elle differe par ses femures anterieurs tres legerement sinues vers leur 
extremite. Elle a aussi una seule Jent devant la sinuosite mediane de femur. 
Un peu plus trapue, de fonne moins allongee que R. jw<ca Pall, de B., ce qui 
pourrait la rapprocher de R. quadridentata Stal, elle se separe aussi franche- 
ment de cette derniere par la forme du pronotum beaucoup plus court. En 
effet I'insecte a 34 mill, de longueur, sur lesquels la tete et le pronotum n'ont 
que 10 mill. Les appendices de 22 mill, sont sensiblement plus courts que 
I'abdomen. Les femurs intermedif»res et posterieurs courts, replies en avant 
depassent a peine la tete. 

Le pronotum tres robuste presque trois fois plus court que Tabdomen, assez 
forttment dilate en a\-ant et tres fortement elargi en arriere, ne permet pas de 
la confondre avec R. kirkaldyi T. B. II est en outre marque de deux sillons 
longitudinaux un peu obliques sus les cotes, derriere la dilatation anterieure, 
n'atteignant pas en avriere les sillons transversaux qui limitent en avant la 
partie posterieure dilatee du pronotum. Cette derniere marquee d'une carene 
longitudinale mediane evanescente en arriere, mieux accentuee en avant ou 
elle traverse les sillons transvei-so^ux qui limitent la partie posterieure dilatee. 

Les pattes pas tres greles, un peu plus courtes proportionnellement que 
celles de R. quadridentata Stal; femurs rougeatres, marques d" anneaux pales, 
larges, peu visibles. 

Metasternum en plaque, termine au milieu en arriere par un prolongement 
retreci entre les hanches posterieures, paraissant plus releve que chez R. fusca 
P. de B. et R. quadridentata Stal, mais moins cependant que chez R. fabricii 
Guer. := annulipes Stal. 

Cette espece se distingue encore des trois autres formes connues de I'Amer- 
ique du Nord par I'opercule genital 9 depassant un peu sous la base des ap- 
pendices. Le segment ventral qui precede I'opercule genital presque droit sur 
son faite longitudinal tres peu convexe avant I'extremite. 

L'espace interoculaire convexe entre les yeux, mais sans trace de tubercule, 
a peine plus large qu'un ceil. Les yeux tres legerement transversaux. 

San Diego, Cal. Coll. Coquillett. U. S. N. M., Washington. 



Hungp:hfori): Nepid.e in America. 449 

Notes. I have examined the type at the United States National 
Museum, the single example from which Doctor Montandon drew 
his description. It is a female. There is also a male with same data, 
that probably was taken with the type, and in addition there are 
two females from Eldorado county, California. One of them bears 
this note: "Fife of this insects were seen feeding on one grass- 
hopper which got into the water." Another specimen, a female, 
from Lindscy, Cal., taken by C. Pemberton, is also in the National 
Museum. I have in my collection some specimens from Laguna 
Beach, Cal., taken by C. T. Dodds. To me the most striking char- 
acter of Ranatra brevicollis is the apparently truncate head; the 
head does not appear broader than anterior part of pronotum, which 
is thick and short. The anterior lateral prolongation of the penulti- 
mate segment of the antenna is short, but little more than half the 
ultimate. 

The salient characters, as assigned to this species by Doctor Mon- 
tandon, are: Body thick-set, pronotum short, appendages shorter 
than abdomen, the legs not very slender, a little shorter proportion- 
ally than those of R. quadridentata Stal; metasternal plate with a 
middle prolongation between the posterior coxae, more elevated than 
in R. fusca P. B. (= R. buenoi, sp. new), and R. quadridentata Stal, 
but less, however, than in R. fabricii Guer. = annulipes Stal. He 
says also that the genital opercule extends a little under the base of 
the appendages and the ventral segment which precedes the genital 
opercule almost straight on its longitudinal summit, very little con- 
vex before the extremity. I find, however, that the male operculum 
does not extend under the base of the appendages. 

All of the insects of this species which I have seen came from 
California. (See fig. 2, pi. XLVIII;'fig. 3, pi. XLV; fig. 14, pi. 
XLVH;fig. 3,pl. L.I 

Ranatra australis, sp. new. 

Size. Smallest specimen in our series measures 32 mm. from tip of beak 
to tip of abdomen with a respiratory tube 27 mm. long. The largest specimen 
is 37 mm. long with a tube 30 mm. long. 

Shape. On the whole a slender species with a long prothorax and long 
hind femora. 

Structural peculiarities. Eyes normal; juga; very prominent, more ele- 
vated than tylus, a characteri.stic that distinguishes this species; antennae 
with lateral prolongation of penultimate .segment nearly as long as ultimate. 
Prothorax slender, sides fairly straight, the anterior portion measured on the 
median dorsal line two more or less times the posterior swollen part. Respira- 
tory filaments quite long, a little less than length of the insect. The dasper 



450 The University Science Bulletin. 

of the male genital capsule with the anteapical prolongation truncate and 
short and well separated from the apical. (See fig. 3, pi. XLVI.) Front 
femora broad without apical tooth. Hind femora surpassing the middle of the 
penultimate segment of the body and the hind tarsus reduced to one-sixth 
of its tibia. (See fig. 4, pi. XLVIII; fig. 1, pi. L; and fig. 15, pi. XLVII.) 

Notes. Described from the following: 

Holotype: Male, Colorado county, Texas, June 24, 1922; Mrs. Grace 

Wiley. 
Allotype: Female, Colorado county, Texas, June 23, 1922; Mrs. Grace 

" Wiley. 
Paratypes: Five males and four females, Colorado county, Texas, June 
23 and June 24, 1922; Mrs. Grace Wiley. 
Nine males and three females, Gainesville, Fla., June, 1918; C. J. Drake. 
One female. New Orleans, La., June, 1915. 
One male, McComb, Miss., July 27, 1921. 
One female, Fayette, Miss., July 23, 1921; C. J. Drake. 
One male. Mound, La.,; J. C. Bradley (teneral specimen). 
One male, Calyell, La., June 16, 1917; H. H. Knight. 
One male, Kissimmee Lake, Florida; A. N.Resse. 

Total of twenty-nine specimens from Texas, Florida, Mississippi 
and Louisiana. The last specimen, belonging to the U. S. National 
Museum, was labeled R. fusca by Doctor Montandon in 1909, and 
bears the following interesting note: "Alligator flea, water dog, said 
to bite or sting severely. Swamp east of Lake Kissimmee, Osceola 
county." 

The holotype and allotype and two paratypes are in the Uni- 
versity of Kansas collection; paratypes are also in the collection 
of Dr. C. J. Drake, Ames, Iowa; the United States National Mu- 
seum, Washington, D. C; J. R. de la Torre Bueno, and the collec- 
tions of Mrs. Grace Wiley and of the author. 

This species is smaller, slenderer and longer limbed than R. fusca 
{=R. americana Montd.). It differs also in the following particu- 
lars : 

1. The jugae more prominent than tylus — not true in R. fiisca P. B. 

2. The eyes are smaller than in R. fusca P. B. 

3. The pronotum is longer; the anterior part two more or less, times the 
posterior part, whereas in R. jusca P. B. it is P/^ to 1%+. 

4. The sides of prothorax more nearly parallel and posterior swollen part 
not so swollen. 

5. The hind margin of pronotum roundly and broadly emarginate, whereas 
in R. fusca P. B. {=zR. americana Montd.) the emargination is deeper and 
narrower. 

6. The two depressions on the scutellum are deep and pitlike, while in R. 
fusca P. B. they are shallow and broad. 



Huncjekfohd: NEPin.ii in America. 451 

7. The hind femora longer, surpassing the middle of the penultimate body 
segment, often almost attaining its caudal margin. Femora not so de- 
veloped in R. fusca. 

8. Hind tarus of R. amtralis one-sixth or less of the tibia; one-fifth or less 
in R. fusca P. R. 

9. Metaxyphus usually longer. 

10. The female operculum angulate on its ventral line, while it slopes gradu- 
ally and is longer in R. jusca P. B. 

11. The respiratory filaments are relatively longer in R. austtralis sp. new, than, 
in R. fusca P. B. 

12. The front femora lack the apical tooth; R. fusca P. B. has one more or 
less marked. 

Ranatra drakei, sp. new. 

Size. Length from tip of beak to tip of abdomen 35 mm. to 46 mm.; iu 
addition to this, the respiratory filaments are from 28 mm. to 44 mm. long. 

Color. All the specimens in the series studied are yellowish brown with legs 
and tegmena overcast with an orange tinge. 

Shape. A long, slender species with prominent eyes; long, slender protho- 
rax; hind femora surpassing the middle of the last abdominal segment, and a 
ver>' long respiratory tube. 

Structural peculiarities. The eyes very prominent, transverse diameter 
greater than interocular space; jugse prominent and divergent; antennae 
with lateral prolongation of penultimate segment a little more than half the 
length of the ultimate segment; prothorax slender, the anterior portion 
measured on the median dorsal line 2V^ times length of the posterior swollen 
part. The length of abdomen is to length of pronotum as 2^/4 is to 1; the 
respiratory filaments long, as long as entire body in many of the specimens, 
greatly surpassing the limbs. The claspers of the male are shown on plate 
XLVI, figure 11. The limbs are long and slender; front fenlora slender, median 
tooth considerably nearer apex than base; distal tooth well marked and 
located at some distance away from the tibial joint, this distance being about 
one-fourth the length of that part of femur lying in front of the median 
tooth (see fig. 3, pi. XLYIII) ; middle and hind femora long; distal end of 
hind femora attaining, or nearly attaining, the caudal margin of the penulti- 
mate abdominal segment; the ratio between femora and tibia not quite but 
nearly as great as in R. buenoi, sp. new. See plate L, photograph 4 of para- 
type specimen.) 

Notes. Described from eleven specimens, seven males and four 
females, taken at Gainesville, Fla., ten of them by Carl Drake, June, 
1918, and one specimen taken March 18, 1915, collector unknown; 
holotype in collection of Carl Drake, allotype in University of Kan- 
sas collection, paratypes in the above collections and in that of the 
author. 

This species has the general appearance of R. buenoi, due to the 
large eyes and elongate, slender body. It differs from that species, 
however, in the front femur possessing a well-defined apical tooth; 



452 The University Science Bulletin. 

in more prominent jugse; in differently formed antennae; in its longer 
thorax; in the respiratory filaments greatly surpassing the limbs 
when extended backwards, and in the differently formed claspers of 
the genital capsule of the male. It cannot be confused with any 
other of our species. 

Ranatra quadridentata Stal. 

Stal, Ofversigt af Kongl. Vetenskaps akademiens forhandlingar, Arg. 18, 18G1, No. 4, 
p. 204. 

Original description: 

Grisea, pedibus immaculatis; abdominis dorso sanguineo; oculis modice 
prominiilis; thorace antice leviter ampliato, subtus bisulcato; alis levissime 
infuscateiA; femoribus anticis subtus pone medium et prope apicem, licet hie 
obsolete, bidentatis, intermediis ]:)osticis vi.x aequilongis, his basim segmenti 
penultimi abdominis vix superantibus; metasterno ut in praecedente. Long. 33 
to 36 milhm. Mexico. (Mus. Hohn.) Prsecedenti affinis. 

The "preceding" is R. unidentata, from Rio Janeiro, and concern- 
ing its metasternum he says, 

"metasterno retrorsum fere ad apicem coxarum posticarum producto, seg- 
mentum ventrale primum tegente, uti videtur postice trilobato, lobis continuis, 
elongatis, medio convexo, lateraUbus depressis, subarcuatis." 

Notes. Van Duzee's catalogue completely submerges United 
States records of this species under R. americana Montd.* This is 
not justified, because Doctor Snow in his list (Trans. Kans. Acad. 
Sci., vol. XX, pt. 1, p. 153, 1906) was not writing about the insect 
that Doctor Montandon described as R. americana. I have before 
me Doctor Snow's insect from San Bernardino Ranch. Cochise 
county, Arizona, the same thing from Mexico, and a series from 
Texas. These insects agree splendidly with Doctor Montandon's 
comparative notes on Stal's species, cotypes of which he has studied 
carefully. 

When Doctor Montandon described his R. americana (Bui. Soc. 
Sci. Bucharest, XIX, p. 65, 1910) he gave some remarkably clear 
notes of comparison between his species and Stal's R. quadridentata. 
The latter is not so robust, the eyes smaller, not so transverse, the 
interocular space not so convex; cheeks not so elongate, but not 
applied so closely against the tylus, which is shorter, making the 
head before the eyes appear shorter. The median construction of 
the pronotum is not nearly so marked and the anterior part more 
cylindrical. I may add that the anterior part of the prothorax is 
longer proportionally, the legs are longer and metaxyphus longer. 

* Van Duzee's catalogue, quadrinotato. Doctor F. H. Snow published a list of the in- 
sects taken by him in Arizona, and listed R. quadridentata Stal as one of them. 



Hungerford: Nepid.e in America. 453 

SUMMARY. 

The Ncpidae of North America north of Mexico now include a 
total of eleven species and two varieties, distributed as follows: 
one Nepa, two Curicta, and eight Eanatra species with two varieties. 
This count omits R. annulipes Stal, which probably does not occur 
in our range. The naming of our two commonest and widespread 
species, under the two oldest names given, is as conclusively shown 
as can.be done without authentic types. We have decided that our 
large, robust, short-limbed form, with broad anterior femora, is R. 
fusca P. B., and that our slender-limbed form, with uniformly short 
respiratory filaments, is R. nigra H. S. This, most unfortunately, 
reduces to synonymy R. americana Montd. and R. protensa Mondt., 
names by which these species have been well known to us for the 
past ten years. The antennae and genital claspers of the males have 
been of value — very striking and satisfactory characters in most of 
the species. The genital capsules themselves are of little value sys- 
tematically in most of the Ranatra. The intromittent organ of the 
male has a more or less definite and characteristic shape at its tip 
(compare figures 6-1 and 8-1, plate XLVIj. Care has been taken 
to examine material from various localities in order to fix the limits 
of variation of these characters. 



454 The University Science Bulletin. 



PLATE XLIV. 

■ Fig. 1. Egg of Ranatra jusca P. B. (=i2. americana Montd.) dissected 
from a water-soaked dead cat-tail blade. 

Fig. 2. Diagrammatic drawing of Ranatra. A, apical tooth of front femur. 
M, median tooth of front femur. C, front coxa. G, prothoracic transverse 
grooves which separate the "anterior part of the pronotum" from the "swollen 
posterior part." S, scutellum. 

Figs. 3 and 5. Stridulating device of Ranatra. Figure 5 shows ventral view 
of head and a portion of prothorax. The rubbing of the roughened patch {A) 
on the base of the coxa (C) against the file (B) on the inside edge of the an- 
terior lateral margin of the prothorax (P) produces the chirping or squeaky 
noise. D, antennae. Figure 3 gives an enlarged lateral view of base of coxa and 
anterior lateral margin of prothorax cleared so that the file shows through. 
Lettering same in both figures. 

Fig. 4. Ranatra eggs in situ in soft, decaying cat-tail leaf, a portion of 
which has been removed to expose the eggs. 

Fig. 6. Genus Curicta. Note that it is intermediate in shape between 
Ranatra and JVepa. Until the appearance of this bulletin only a single speci- 
men was recorded from the United States. 

Fig. 7. Egg of Curicta drakei, sp. new, dissected from the tissues of a de- 
caying plant stem, where only the crown of fifteen filaments was exposed. 
Drawing made from material secured by Mrs. Grace Wiley in her life history 
notes on this species. 

Fig. 8. Egg of Nepa apiculata Uhl. Note the eleven filaments which remain 
exposed above the surface of the plant in which the eggs are inserted. 

Fig. 9. Genus Nepa. Represented in the United States by a single species, 
Nepa apiculata Uhl. This species varies considerably in size, but no constant 
structural detail has been found to indicate that the variants are not conspe- 
cific. 



Hun'gerford: Nepid.e ix America. 



455 



PLATE XLIV. 




456 The University Science Bulletin. 



PLATE XLV. 

Fig. 1. Clasper of male of R. elongata Fab., determined by Doctor Mon- 
tandon. S, subapical tooth. 

Fig. 2. Genital capsule of male of R. elongata Fab., determined by Doctor 
Montandon. Compare figures 1 and 2 with figures 3 and 4. Note the differ- 
ences in shape of claspers, their relative lengths as shown in figures 2 and 4. 
The intromittent organ, I, is also often characteristic of a species, but not 
always, and subject to some variation. The capsule itself in Ranatra is of 
little value. 

FiGS^ 3 AND 4. Clasper and capsule of male R. brevicollis Montd. from 
California. 

Fig. 5. Inside view of left clasper of Curicta drakei, sp. new. Compare 
with figure 8, C. howardii Montd. The shape of the clasper near the base is 
not the same in the two species. 

Fig. 6. Male genital capsule of C. drakei, sp. new, viewed from right side. 

Fig. 7. Male genital capsule of C. howardii Montd. 

Fig. 8. Inside view of left clasper of C. hoxmrdii Montd. 

Fig. 9. Ventral view of head of Ranatra to show the antennae, which lie 
hidden. Sometimes they are covered with debris, which should be scraped 
away. It is often wise to relax the specimen and draw the antenna into a 
more exposed position. The front coxa also sometimes obscin'es the view and 
should be moved while specimen is relaxed. 

Fig. 10. Lateral view of caudal end of abdomen and base of respiratory 
filaments of Ranatra nigra H. S. C, connexivum. R, respiratorj' filaments. 
0, operculum of genital segment, considered in this paper as last abdominal 
segment. P, considered in this paper as the penultimate abdominal segment. 

Fig. 11. Lateral view of caudal end of abdomen and base of resjiiratory 
filaments of Ranatra jabricii Guer. from Cuba. Montandon says it is identical 
with R. annulipes Stal from Brazil. 

Fig. 12. Dorsal view of head of Ranntra. B, beak. ./. jugum. T. tylus. 



Huxgekfukd: Nepid.k in America. 



457 



PLATE XLV. 









458 The University Science Bulletin. 



PLATE XLVI. 

Figs. 1 and 2. Clasper and male genital capsule of Ranatra buenol, sp. 
new. Note the reduced subapical tooth of the clasper and the spatulate form 
of the tip. 

Figs. 3 and 4. Ranatra australis, sp. new. Note the truncate subapical 
tooth of the clasper and its distance from the apical one. 

Figs. 5 and 6. Ranatiia kirkaldyi Bueno. S, subapical tooth. C, clasper. 
I, intromittent organ. This species is clearly quite distinct in its relationship 
from the others. Note the shape of the intromittent organ and of the clasper. 

Figs. 7 and 8. Ranatra quadridentata Stal. I am not able to find any 
constant differences between this and R. jusca P. B. {^R. americana Montd.) 
in respect to the male genitalia. The male of the latter species has, on the 
whole, a more slender clasper. 

Figs. 9 and 10. Ranatra brachyura Horv., 1879, said to be same as R. 
sordidula Dohrn., 1860. From Japan. Confused by a student of this family 
with R. protensa Montd. and so labeled. Compare figures 9 and 13. 

Figs. 11 and 12. Ranatra drakei, sp. new. 

Figs. 13 and 14. Ranatra nigra H. S. (^R. protensa Montd.). The pre- 
ceding species and this one have claspers which are broad in their middle 
parts, but the shape at the tip distinguishes them. 

Figs. 15 and 16. Ranatra linearis Fab. From Euroj^e. Berlese. in his 
Gli Insetti, p. 323, figures this species. Note that the difference between the 
clasper of this species and that of our American species is more apparent than 
the superficial appearance of the insects when side by side in the cabinet. 



Hungerford: Nepid^e in America. 



459 



PLATE XLVI. 













460 The University Science Bulletin. 



PLATE XLVII. 

Fig. 1. Ventral view of male genital capsule of Nepa apiculata Uhler. 
J'rom North America. 

Fig. 2. Lateral view of clasper of above. Compare with figure 4 of N. 
cinerea L. of Europe. 

Fig. 3. Ventral view of male genital capsule of Ncpa cinerea L. 

Fig. 4. Clasper, lateral view of Nepa cinerea L. 

Fig. 5. Antenna of Nepa apiculata Uhler. From New York. 

Fig. 6. Antenna of Nepa cinerea L. From Europe. U, ultimate antennal 
segment. P, penultimate antennal segment. L, lateral prolongation of penulti- 
mate segment. The splendid antennal and genital characters separating these 
two species of Nepa illustrate the taxonopic value of these hitherto unused 
structures in the Nepidse. In the drawings which follow, do not attach sig- 
nificance to the basal segment, but to the last two segments. 

Fig. 7. Antenna of Curicta drakei, sp. new. 

Fig. 8. Antenna of Curicta howardii Montd. 

Figs. 9 .and U. Antennse of Rnnatra kirkaldyi Bueno. Sometimes the ulti- 
mate segment is fused with the preceding segment and sometimes entirely 
separate. 

Fig. 10. Antenna of Ranatra nigra H. S. (^R. protensa Montd.). 

Fig. 12. Antenna of Ranatra buenoi, sp. new. 

Fig. 13. Antenna of Ranatra drakei, sp. new. 

Fig. 14. Antenna of Ranatra brevicollis Montd. 

Fig. 15. Antenna of Ranatra australis, sp. new. 

Fig. 16. Antenna of Ranatra quadridentata Stal. 

Fig. 17. Antenna of Ranatra fiisca P. B. (=J?. americana Montd.). 



HuNGERFORn: Nepid.k in America. 



461 



PLATE XLVII. 








II 







462 The University Science Bulletin. 



PLATE XLVIII. 

Fig. 1. Front femur, tibia and tarsus of Ranatra buenai, sp. new. Note its 
very slender form and the position of the median tooth. 

Fig. 2. Ranatra brcvicollis Montd. The median tooth is nearer the middle 
than in any of the others. The tibia is relatively longer. 

Fig. 3. Ranatra drakei, sp. new. Note the slender form and the position of 
the aincal tooth. 

Fig. 4. Ranatra australis, sp. new. It lacks the apical tooth, and the an- 
terior portion of the femur is not enlarged toward the distal end as in 
R. jusca P. B. 

Fig. 5. Ranatra fiisca P. B. (= R. ainericana Montd.). From Ithaca, N. Y. 

Fig. 6. Ventral view of head and prothorax of R. buenoi, sp. new. To show 
the deep longitudinal trough. Compare with figure 10, which lacks it. 

Fig. 7. Ranatra kirkaldyi Bueno. Note that the femur is not constricted 
in the region of the median tooth. Compare with R. fusca P. B., figure 5. 
This from Kansas. 

Fig. 8. Ranatra nigra H. S. {=: R. protcnsa Montd.) 

Fig. 9. Ranatra kirkaldyi Bueno var. hofjnianni new. Apical tooth or 
marked sinuosity present. From Minnesota. 



Hiingerford: Nepidk ix America. 



463 



PLATE XLVIIT. 




464 The . University Science Bulletin. 



PLATE XLIX. 

(All photographs natural size.) 

Photo 1. Photograph of Palisot's figure of Ranatra jusca, reproduced exact 
size. 

Photo 2. Ranatra kirkaldyi Bueno. Largest specimen in a long series. 
A female from Cherokee county, Kansas, determined by Mr. J. R. de la 
Torre Bueno. This is what Doctor Montandon has mistaken for Ranatra jusca 
P. B. Besides size, note shape of front femur. Not constricted as it is in 
figure 1. 

Photo 3. Ranatra jusca P.B. (= R. americana Montd.) Note the general 
resemblance to original figure by Palisot (photo 1 above), the shape of the 
thorax, the broad anterior femora, the size of the insect, the reduced hind 
femora, etc. For other figures of this insect, see plate LI, photographs 3 and 7. 
Figure 7 is the only one to show the apical tooth of the front femur plainly. 
The writer has forms with the respiratory tube of various lengths. The short- 
est perfectly formed tube is shown on plate LI, figure 3. 

Photo 4. Ranatra kirkaldye Bueno var. hojjmanni, var. new. 

Photo 5. Ranatra bitenoi, sp. new {=R. jusca Bueno). Compare with 
Palisot's R. jusca (photo 1). The relative proportions of the front femora, the 
length and shape of thorax, limbs and filaments. The long tibise and reduced 
tarsi. This photograph is of a paratype. 



PT.ATI-: XT.TX. 





\\ 




(465) 



PLATE L. 

. (All photographs natural size.) 

Photo 1. Ranatra australis, sp. new. Differs from R. jusca P.B. by shape 
of jirothorax. longer respiratory tube, lack of apical tooth on front femora 
and by its elevated jugse, as well as marked differences in the genital claspers 
of the male. Southern in distribution. Photograph from paratype. 

Photo 2. Ranatra nigra H. S. (= R. protcnsa Montd.) This insect in a long 
series fits the original description most remarkably, except for the color, which 
is of no consequence at all; "Size 2-2 Vs inches from beak to tip of filaments. 
Filaments % inch long; limbs long and slender." Photo 5 is another speci- 
men — the latter from Minnesota, the former from Kansas. 

Photo 3. Ranatra brevicollis Montd. From California. Note the short, 
broad thorax and its relation to the size of the head. 

Photo 4. Ranatra drakei, sp. new. Slender front femora with apical tooth, 
large eyes, long thorax, long limbs, and Aery long respiratory filaments. 
(Photograph from paratype.) 

Photo 5. Ranatra nigra H. S. (^R. proteiisa Montd.) 

(466) 



PLATE L. 





/ 



J 



/ 



(467) 



PLATE LI. 

(All photographs natural size.) 

Photo 1. R. quadridenlata Stal. Small specimen taken by Doctor Snow- 
in Arizona. 

Photo 2. Curicta howardii Moutcl. Photograph from male belonging to 
Professor Barber. Compared with type at Washington by Doctor Drake. 

Photo 3. Ranatra fusca P.B. Specimen from Minnesota with very short 
but perfectly formed re.spiratory filaments. The tijjs are normal. 

Photo 4. Curicta drakei, sp. new. Male holotype. 

Photo 5. Nepa cinerea L. Male from Europe. Compare with A^cpa api- 
culcUa Uhl. from Minnesota, photo 8. then note structural differences figured 
on plate XLVII. 

Photo 6. Curicta drakci, sp. new. Female allotype. 

Photo 7. Ranatra fusca P.B. Shows how long the respiratory filaments 
may be and how marked the apical tooth of front femur. 

Photo 8. Nepa apiculata Uhl. Female from Minnesota. 



(468) 



PLATE LI. 





/ 




/ 







8 



(469) 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 19— October, 1922. 

(Whole Series, Vol. XXIV, No. 19.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

A Study of the Relation Between Function and Growth 
IN Body Cells Montrose T. Burrows. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second-class matter. 

0-4622 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV.] October, 1922. [No. 19. 



A Study of the Relation Between Function and Growth 

in Body Cells. 

BY MONTROSE T. BURROWS, M. D. 

The Department of Surgery, Washington University School of Medicine and the Barnard Free 

Skin and Cancer Hospital, St. Louis, Mo. 

IN THE BODY, growth is a definitely regulated act, which dur- 
ing development takes place irregularly in one part and then in 
another. At maturity it ceases in most tissues, except as it is neces- 
sary to replace tissue or cells lost by injury, to meet the demands 
of an increased function of the whole or a part, and to take care of 
the general wear and tear. Function, on the other hand, such as 
heart-muscle contraction, is something that, once established, goes 
on continuously throughout the life of the individual. 

Growth in the body is, therefore, that which has a maturity or 
has limits, while function (rhythmical heart-muscle contraction) 
has none. 

These two processes are related in only one regard: for a given 
amount of work of contraction there is a constant size. A stimulus 
which induces an increase in work on the part of the heart induces 
also an increase in the size of the organ, while one which induces a 
decrease in work leads to atrophy. What is true for the heart is also 
true for muscles in general and for the kidneys, liver and other or- 
gans of the body. Growth of the muscle fibers and cells of these or- 
gans is proportional, therefore, to their rate of activity or to the rate 
of their metabolism. That the same is true in the embryo, even be- 
fore differentiation and the development of function, has been clearly 
pointed out by Child.* ^ Child again points out that with each de- 
crease in growth rate there is a decrease in the rate of metabolism, 

* Superior figures refer to numbered paragraphs of the bibliography at the end of this 
article. 

(475) 



476 The University Science Bulletin. 

and that this metabolic rate increases markedly in lower forms after 
starvation or when they are regenerating lost parts. 

The question of growth regulation may, therefore, be temporarily 
set aside for a study of the conditions which regulate metabolism in 
general in the cell. As the tables for growth clearly show, the 
metabolic rate in the organism is never constant (Child and Thomp- 
son).^ In early embryonic life the increment is high, then it gradu- 
ally decreases, with irregularities, up to the death of the individual. 
In the early stages of development the blastomeres are simple, ex- 
panding, growing cells. Later they differentiate and assume a func- 
tional state. That this differentiation and the development of func- 
tion is brought about by the continuous changing environment ef- 
fected by the growth and filling up of cells in the growing individual 
has been clearly indicated by many already well-known studies of 
experimental embryology.^ What we have not known are (1) the 
nature of those conditions which bring about these changes, and (2) 
the importance of these changes for the life of the organism itself. 

In later life we have learned to recognize the fact, however, that 
all tissues do not become actively functioning cells. A part of the 
mesenchyme changes to muscle and glands, while another part re- 
mains in what appears to be a more primitive state, such as the cells 
of the interstitial tissue, bone and cartilage. The same is true for 
epithelium. A part is used for covering and protection and another 
part becomes functioning glands. For the sake of simplicity, I shall 
call the cells w^iich are active in the glands, in the muscle, etc., as 
functioning cells, in contradistinction to those of the connective tis- 
sue, bone, cartilage and the epithelial coverings, the nonfunctioning 
cells. This is not a strict use of the terms, but the latter resemble 
more closely in their behavior the undifferentiated cells of early life. 

There was a time when it was thought that all life processes cen- 
tered within the cell. At the present time there is some question 
whether this is true. The primary changes in early development ap- 
pear to be a rapid swelling of the mass, a rapid inhibition of water 
and oxygen, a rapid elimination of CO2, and a secondary splitting 
of the whole into cells. Cellular growth, division and differentiation 
are not primary activities, but apparently secondary to other more 
formative forces or stimuli. Life does not manifest itself the same 
throughout the life of the animal. In the beginning it is recognized 
chiefly by the rapid expansion of the whole, the rapid division of 
cells and the careful building of its various parts. This period of 
building is completed in man, as is well known, within ten days after 



Burrows: Study of Body Cells. 477 

birth. Subsequent to this time no new oi-^an or parts of organs are 
formed except the hiying down of the nerve sheaths. All subse- 
quent growth is merely the expansion of previously formed struc- 
tures. It is like hypertropliy and hyperplasia, as they result from 
stimulation of the adult organs. 

With the completion of the building of the organs, function has 
made its appearance. Growth unlike that of the earlier period now 
runs hand in hand with new forms of work. This growth continues 
to maturity, when it ceases, except as it is to take care of the wear 
and tear, so to speak, and to play a part in certain organs and tis- 
sues, such as the bone marrow, the sex glands, and the nail- and 
hair-beds. Atrophy then slowly intervenes. This continues to the 
inevitable death. 

The general nature of the structure and the metabolism which 
leads to this sequence of changes in the organism is the pertinent 
biological question to-day. ChikU in 1915 reviews the general the- 
ories that have been advanced and attempts to give a physicochemi- 
cal explanation of the process. Child, appreciating the relation 
between the slowing of growth and the decrease in metabolism, 
makes the general assumption, applying the "law of mass action," 
that the decrease in metabolism may be explained as the result of 
the gradual accumulation within the cell of an insoluble substratum. 
Since differentiation and the development of function takes place 
hand in hand with this slowing of metabolism, he looks upon them 
also in the same light. He considers the changes in the structure — 
differentiation — the result of the same slow deposit of the same sub- 
stratum. This substratum he conceives as one of the products of 
the metabolic reaction. The reaction then, like any incomplete re- 
versible physicochemical reaction, becomes slowed and this sub- 
stratum accumulates, and ceases when it reaches a certain con- 
centration. Death is the equilibrium point for this reaction. 

While there is little doubt that many heterogeneous, like many 
homogeneous, reacting systems reach their equilibrium by this route, 
there does not seem to be sufficient evidence to show that the body is 
exactly of this kind. Again, all such systems which do arrive at 
such an equilibrium are first put together by some external force. 
In nature one cannot obtain more energy from any system than what 
has been put into it. This is the law of the conservation of energy. 
.Child looks upon rejuvenation, then, as the result of the removal of 
this substratum and of dedifferentiation. Differentiation is not, there- 
fore, according to Child, a change peculiar to a definite period, but 



478 The University Science Bulletin. 

rather the result of growth, or the dynamic state of the cell. Senes- 
cence is an inevitable consequence suffered by all cells, whether they 
be unicellular organisms or cells of the metazoa. To put it in his 
own words, "Senescence is a necessary and inevitable feature of 
growth and differentiation, while rejuvenescence is associated with 
reduction." Differentiation he defines as a process of specialization. 
It takes place in the cells when they are suffering a moderately ac- 
tive metabolic rate. Decrease in this rate favors differentiation. 
Increase it and dedifferentiation tends to occur. Reduction is dif- 
ferent. It is not the reverse of growth. Growth is the "accumula- 
tion of certain substances formed in the course of the reaction which 
are physiologically more stable than other substances that break 
down, furnish energy, and are eliminated." Reduction takes place 
when the breakdown is not balanced by the synthesis. Using 
Child's own data, I see no reason why this process may not be 
formulated, however, in a much more definite manner and in a man- 
ner more consistent with not only other natural phenomena, but also 
with the picture of development and later life as they have pre- 
sented themselves through morphological studies. In the early 
periods of development, just after the fertilization of the egg, the 
rate of metabolism is high. The work performed during this period 
is the building of a heterogeneous mass out of a previous simple egg 
cell. This process of building is completed early. The metabolism 
at the time of completion is lower than at previous times, but it is 
still high. Subsequent to this time this metabolic rate, then, de- 
clines progressively to death. The picture at the beginning is there- 
fore entirely different from the later one. It is the building of the 
machine, which, when once established, slowly runs down. The pic- 
ture in development is that of the forcible putting together of parts, 
which are then to react with one another to produce work. This re- 
action continues in each case to an equilibrium or death, like any 
such machine built by man. While the metabolic rate is high in this 
machine it grows. This growth ceases (maturity) when this meta- 
bolic rate reaches a certain minimum. Then atrophy slowly sets in. 
Normal death is not the accumulation, but the gradual using up of 
the parts. It is the passing of heterogeneous system to a state of 
equilibrium. 

Such an explanation looks upon the life of the organism, there- 
fore, as the result of some unknown force, active only in its early 
period, beginning with the fertilization and ending with the forma- 
tion of the last organ and parts of organs. This force disappears at 



Burrows: Study of Body Cells. 479 

birth or thereabouts. Subsequent to this time hfe is merely an ex- 
pression of the gra(hial deterioration of the previously built hetero- 
geneous system or the interaction of its various parts, which follow 
tlie law for the disintegration of all such heterogeneous systems. 
Life of the animal ends with the establishment of an equilibrium be- 
tween the parts of the organization which produce the work that 
really constitutes life. The energy for the building of the first 
period is derived, therefore, in the later period. 

Systemic or general death is nothing more, therefore, than the 
establishment of a true equilibrium, or a breakdown in one of the 
essential parts of the body. This is not the end, however. What 
we have been talking about so far is the life of the organism, and 
not that of the cell. The end of the individual does not mean the 
end of all its parts. The cells are not dead at this period. They are 
all intact. This system is able to reproduce itself. The normal 
death of the whole is not the result of the death of the cells. The 
cells which make up the whole are destroyed, rather, by the process 
of the death of the whole. These cells, as we now know, go on for- 
ever under the proper circumstances. Out of one of them in the 
old, the egg cell, comes a new individual. Their destruction at the 
death of the individual is the result of this general death. Their 
destruction is the result alone of their position. Systemic life and 
systemic death are something different, therefore, from elemental 
life and elemental death. The first has limits; the other may have 
none. The question is. What is the nature of this system which can 
show such changes? This cannot be solved by a study of the animal 
as a whole. It cannot be solved by the study of the amoeba, be- 
cause it is an animal itself. It must be solved through a study of 
those cells which go through these changes, those cells which find 
this building of an animal and the ultimate disintegration of this 
animal their normal means for preserving their kind. The amoeba 
need take no such complicated route to preserve its kind. These 
cells of the animal must take this course in order that they survive 
and that the proper environment be prepared for their reproduction. 
Child has ignored this fact completely. He has assumed a continu- 
ous dynamic state for the cell without any proof of its existence. 
He has again assumed that the life of the whole is only an elabora- 
tion of the life of the cell. He has assumed differentiation to be a 
peculiarity of all cellular life. He has assumed the protoplasm of 
cell to have structure capable of performing work. The best bio- 
logical work of the last century has not only added no proof for 



480 The University Science Bulletin. 

these general assumptions formulated by the earlier authors, but 
quite the reverse, it has spoken against them. As Bayliss states, 
there is not evidence that cells are necessarily dynamic. The best 
morphological studies of protoplasm have again failed to show struc- 
ture in many cells other than nucleus and centrosome. As Wilson 
clearly states, cellular growth, division and differentiation are not 
primary factors in development, but secondary to more formative 
forces or stimuli. In the simple formulation which I have given above 
it is possible to understand how the organism not only reproduces 
itself, but it expresses definite need for the differential changes as 
they come into existence. The energy for the primary building is 
acquired from the old. The old is a machine not different in prin- 
ciple from other machines of nature. 

As I shall show in the following pages, the main criticism of 
Child's theory is that he utilizes theories to build theories. He ac- 
cepts the idea of the cell as the unit of manifested life of the organ- 
ism without question. Before any theory of life and death is justi- 
fiable of acceptance it is necessary that the true nature of the struc- 
ture and the metabolism of the cell be ascertained. This is not go- 
ing to be accomplished by morphological and chemical methods 
alone, nor by a study of the metabolism of the whole, but by 
methods which allow us to study directly each of the fundamental 
manifestations of life, such as growth, division, differentiation, 
migratory movement, etc. The tissue culture has given us this 
opportunity. In support of the above contentions it is of interest to 
report here some general analyses so far carried out by this method. 

THE CONNECTIVE-TISSUE CELLS. 

As is well known, practically all previous work on the cell in re- 
gard to the nature of its energy-producing reactions and the man- 
ner of the transformation of this energy has been based upon the 
idea that the cell is a highly organized body. While for years it has 
been assumed that amoeboid movements are comparable to surface 
tension changes in liquids, all theories have been based upon the 
idea that these amcsboid movements are the result of localized 
changes in surface tension resulting from some unknown organiza- 
tion residing within the cell. All modern physiochemical methods 
have failed, however, to reveal any such organization. This has led 
one and then another to assume that the organization is either the 
result of a slow diffusion of substance in the colloids of the cell 
(Wells) ^, a peculiarity of colloids not yet organized, or to invisil'le 
membranes traversing the protoplasm. 



BiRRows: Sttdy of Body Cells. 481 

As early as 1913" I had noted, however, that the movement of the 
cell of the organism is not one which is governed necessarily by such 
factors.^ These cells in the medium of the cultures move always 
out and away from other cells ; and in further studies carried on over 
several years, I have continually noted that the large number of 
these cells show no change in contour during their movement. Their 
movements are not amoeboid. They glide along like bodies carried 
by some external force. 

The picture observed in these cells is not that of a highly organized 
body, but one liberating a surface-tension-lowering substance. 
Their movements are like those of cayenne-pepper granules dropped 
on the surface of water. When a number of such granules are scat- 
tered on the surface of the water they shoot apart. This moving 
apart is the result of the liberation by them of a surface-tension- 
lowering substance. This accumulates in greater concentration be- 
tween the granules. These granules are pulled apart, therefore, by 
the greater force of the water surface without.'' 

While the connective-tissue cells take the same course outward, 
they never become completely dispersed. These cells again fail to 
show movement on the surface or within a liquid medium, but move 
only and show evidence of metabolism in the presence of the fibrino- 
gen contained in the blood plasma which I had used chiefly as a 
medium. In a liquid medium these cells round off to perfect spheres. 
In the plasma cultures they stick tightly to the fibrin formed in the 
coagulation of the plasma. In contact with these fibrils they spread 
out to take various shapes. These shapes are always peculiar to the 
surface of their contacts.^" 

Not only the character of the movement of these cells, but their 
general effect upon the clot, further indicates that they liberate such 
a surface-tension-lowering substance. This substance differs in its 
physical properties from that liberated by the granules of cayenne 
pepper, however, in that it is apparently not soluble in water, but it 
is adsorbed or chemically combined with the fibrinogen. The cells 
not only stick tightly to the fibrin, but they occasion its formation. 
When a fragment of connective tissue is placed in a drop of plasma 
it occasions first a gelation of the whole of the layer, and then later a 
true coagulation. This gelation commences at the tissue border and 
spreads rapidly, to invade ciuickly the whole of a large area of the 
fluid plasma. After a considerable latent period, the coagulation, the 
formation of fibrin and serum, commences. This true coagulation 
again begins at the tissue border and spreads slowly outwards, to 



482 ' The University Science Bulletin. 

involve after many days a small area about the fragment.^^ With 
this second coagulation the cells appear. They are in close contact 
with the fibrin and they glide out just behind the spreading area of 
change in the jelly like clot. This movement continues only so long 
as the process of coagulation proceeds. The cells are elongated 
spindles closely cemented to the fibrin, yet capable of gliding on its 
surface. With the completion of the coagulation their movement 
ceases. They come to rest. In this state they will remain ap- 
parently indefinitely unless fresh fibrinogen is added. When this is 
added they again occasion its coagulation, move into it, and again 
come to rest when the coagulation is completed. 

The whole picture of activity in these cells is that which can be 
readily interpreted in terms of the liberation by them of some sub- 
stance insoluble in body fluids, but readily adsorbed or chem- 
ically combined with fibrinogen. The combination leads to the 
formation of fibrin. The movement and the clinging of the cell to 
the fibrin indicates further that this substance is one which has 
strong affinities for the cell. These cells do not crawl, but they 
glide, and are held firmly to the fibrin. Their gliding is directly 
proportional to the spread of the coagulant. Such can take place 
only in presence of a substance which is strongly attracted, not only 
by the fibrinogen, but also by the cell. Energy production in these 
cells is centered, therefore, about this substance. How it is formed in 
the cells becomes, then, a problem of interest. The fact that the 
cells come to what appears to be complete rest in the coagulated 
clot indicates that it is the product of the one reaction taking place 
within them. 

For several years it has been evident that energy production in the 
body is the result of chemical change. It is further recognized, as 
Bayliss*^ clearly points out, that hydrogen and carbon enter chiefly 
into this reaction. Nitrogen in the body is used largely for the build- 
ing of the substratum, proteins, in which these reactions proceed. 
This indicates that the reactions leading to protein syntheses are 
different from those producing energy. 

What has not been shown is the nature of the products formed 
in this energy-producing reaction. It has been thought that oxygen 
enters directly into this reaction, and at one time it was thought 
that the products formed were largely carbon dioxide and water. 
Recent careful experiments have indicated that this is not true. 
Fletcher'- has shown clearly that oxygen does not enter into the 
contraction phase of muscle, but is adsorbed largely in the recovery 



BiRROAvs: Study of Body Cells. 483 

pcriocl. Muscle may contract for a time without oxygen. J. Loeb^^ 
has found that oxygen is nceessary to preserve «the structure of the 
fertilized egg. These experiments of J. Loeb also indicate that the 
metabolic reaction may proceed without oxygen. That the same 
is true for the animal colls has long been known by pathologists. 
Cut off the blood supply from any part and it undergoes a coagu- 
lative necrosis. These observations suggest strongly, therefore, 
that oxygen plays a necessary but probably a secondary role in this 
reaction. It acts to remove certain products of the primary reaction 
rather than enter into it. 

All chemical reactions, as it is now fully appreciated, are governed 

not only by the concentration of reacting substances, but also by the 

concentration of one or all of the products formed. Equilibrium for 

a simple reversible reaction is expressed in the following formula: 

A + B + C + . . . = A'4-B' + C' + . . . 

In the animal organism growth is not determined by food any 
more than it is determined by oxygen, but by other unknown factors. 
Bardeen showed that planaria regenerate their parts when starved as 
when they are fed.^"* Morgan found that salamanders regenerate 
their legs as rapidly when starved as when fed.^^ The difference is 
tiiat the starved animals suffer marked emaciation and a general 
atrophy of their organs. 

The above observations of the connective-tissue cells indicates 
clearly the existence of a previously unrecognized product of cell 
metabolism which is evidently insoluble in liquid medium, but 
rapidly adsorbed and chemically combined with fibrinogen. This 
substance, which I shall designate as "L," is also an active blood 
coagulant. It is something, therefore, which is insoluble in water, 
but capable of combining with fibrinogen to form an insoluble com- 
bination, fibrin. It acts in every regard, therefore, like the lipoid 
fraction of the blood coagulant recognized by Woolbridge^^ and 
recently carefully studied by Mills. ^'^ This fraction is a phospholipin 
or a group of such bodies. 

In a recent series of experiments^ I undertook, therefore, to 
ascertain more carefully the exact relation to this substance of the 
general chemical changes of these cells. These cells had apparently 
become inactive after they had come to rest in the clotted fibrin. 
They had laid for months in this inactive state and in the presence of 
oxygen. When removed to fresh medium they had again become 
actively migrating cells. I removed the oxygen from about these 
cells during their movement and also after they had come to rest. 



484 The University Science Bulletin. 

While they are migrating and the coaguhition is taking place they 
disintegrate in the absence of oxygen. They suffer a coagulative 
necrosis. After they come to rest they suffer no change in the 
absence of gas. In the same way, these cells suspended in* a liquid 
are unaffected by the absence of oxygen. In a liquid they do not 
metabolize any more than in the presence of a fully formed fibrin. 
Any slight stimulus which occasions their movement occasions also 
their disintegration in an atmosphere of nitrogen. 

There seemed little doubt, therefore, that in the identification of 
this "L" substance the regulator of the metabolism of these cells 
had been found. It is not something which is continuously washed 
away by the blood stream, but, quite to the contrary, it has very 
specific affinities, -and when brought into contact with fibrinogen it 
forms an insoluble fibrin. This suggested strongly, therefore, that 
continuous activity in these cells must depend not alone on food and 
oxygen, but also upon very special conditions for removing these 
primary products from the cell.^ The only plentiful substance 
which I have been able to find in the adult body capable of such 
removal is fibrinogen. This is transferred by the "L" substance 
into fibrin. In the body the messenchyme cells of older embryos 
and the connective-tissue cells of adults lay down extracellular 
fibrils. Hertzler^^ has shown very definitely that fibrin forms the 
basic proteins of these extracellular or collagen fibrils. ^^ These 
results of Hertzler have been confirmed by Baitsell-^ and myself.^^ 

These observations indicate that the connective-tissue cells of the 
body are not in any sense continuously active. Their only function 
is the production of extracellular fibrils. They do not secrete these 
fibers, but coagulate certain proteins formed elsewhere in the organ- 
ism. The increase in these fibrils is alone indicative of an active 
metabolism in them. The dynamic state of the organism is in no 
sense, therefore, an elaboration of the dynamic state of these cells. 
It must depend on other conditions. This led me to investigate more 
carefully the process of rhythmical contraction as it is seen in heart- 
muscle cells. 

In my earlier studies of the tissue culture I had already shown 
that differentiation in heart muscle is a purely reversible phenome- 
non.^^ The actively contracting heart-muscle cells are derived pri- 
marily from the undifferentiated mesenchyme. When the fragments 
of this tissue are brought into contact with the plasmatic medium of 
the culture the heart-muscle cells at the edge of the fragment which 
had been contracting lose this property at once. They migrate into 



Burrows: Study of Body Cells. 485 

the medium and behave exactly hke the simple undifferentiated 
mesenchyme cells. In 1912 I ^^ showed, however, that in a few 
cases these cells which migrate into the medium may differentiate 
again and develop rhythm. When this takes place it is interesting 
to note that the cells by chance alone had come into a very peculiar 
relation with the medium, and again it is interesting to note that this 
rhythm never develops while the cells are migrating, but always 
after such migration has ceased and the coagulation process is com- 
plete. 

The differentiation of the heart-muscle cells in the outer medium 
takes place very infrequently. In the large majority of the culture 
of the older embryos and adults the clots cling tightly to the frag- 
ment. The cells migrate in contact with the surface of the fibrin 
fibrils. When differentiation takes place the process is different. 
The clot loosens and contracts away in mass. If the ends of a few 
cells remain attached to this clot they may become stretched 
through the serum cavity between the surface of the medium and 
the end of the fibrin fibrils, or between the fragments and the end of 
these fibrils. After the coagulation is complete, these cells, and 
these cells alone, develop rhythm. If they be removed from those 
contacts and be placed in the outer medium in contact with the 
surface of the fibrin they stretch out again and behave like simple 
mesenchyme or connective tissue cells.^ 

By these observations it became possible, therefore, to clear up 
the difficulty of the earlier observations of the connective-tissue 
cells. Dynamic states in the organism, such as the heartbeat, are 
not a property of the cell, but that of a peculiar organization of the 
environment. These cells may produce the energy for the work of 
the body, but there is no evidence that the transformation of this 
energy into work is the product of a cell organization in the case of 
the heartbeat any more than it had been found to be in the case of 
cellular migration. 

What is true, for the connective-tissue cells I find also to be true 
for the epithelial cells of the skin and many of the glands. The 
gland cells in the cultures lose the form peculiar to them in the or- 
ganism. They stretch out, like the skin epithelium, to form broad, 
thin sheets of cells. These cells cannot metabolize except in the 
presence of fibrinogen or a similar adsorbing substance. They differ 
from the connective tissue in that they later destroy this substance 
through certain added proteolytic properties. They thus depend 
wholly on the fibrinogen for their activity in the culture, but they 
remain together and form no extracellular fibrils, in that they later 



486 The University Science Bulletin. 

destroy the fibrin which is formed.-^ In Hquid media they show no 
evident activity. 

While these experiments are interesting for the understanding of 
the general problems of the genesis of connective tissue, their 
broader significance lies in the new view they present of life in the 
organism. They show that the continuous activity consistent with 
life is not a property of these nonfunctioning cells. There must be 
long periods of time when the connective-tissue cells show no ac- 
tivity. Life must be, therefore, wholly a part of the functioning 
systems, the glands, the muscles, the brains, etc. In early life, how- 
ever, this cannot be true. The whole problem of life in the early 
embryo is centered about an excessive metabolism and an active 
proliferation of the undifferentiated cells of this earlier period. 

Many years ago Hofmeister, Sachs and De Bary had already 
shown that cellular proliferation is not a primary factor in the 
growth of plants. They thought that the mass forms in growing 
plants before it breaks into cells. That the same is true for animals 
has been clearly enunciated by Whitman, Adam Sedgwick, E. B. 
Wilson and others. In development, cellular growth, division and 
differentiation are not primary factors, but they are again secondary 
to other more formative forces. Driesch looked upon this force as 
something apart from nature which cannot be solved. Others have 
not looked at it in this sense. They have considered that this early 
development is the result of certain reactions which occasion such 
a primary building. Many general physiologists have attempted to 
arrive at its solution through the study of colloidal swelling. It is 
evident, however, that such study cannot attain directly to this goal. 
At the best they can only indicate analogies. It occurred to me to 
attempt the solution by another method. The above studies were 
made with connective-tissue cells of adults and the mesenchyme 
cells and heart-muscle cells of older embryos. No careful compari- 
son had been made of these cells with those of the younger embryos. 
In the younger embryos it is well known that the mesenchyme cells 
form no extracellular fibrils, but grow actively to form a cellular 
syncytium. The fibrils appear only in later embryonic and adult 
life. 

THE BEHAVIOR OF THE CELLS OF THE YOUNGER EMBRYOS. 

As I have cited in a previous article, M. R. and W. H. Lewis, in 
1911,^^ noted an active movement and growth of cells in liquid 
medium such as simple salt solution. This, as I have just stated, is 
not true of the adult connective-tissue cells. Harrison^^ in analyzing 



Burrows: Study of Body Cells. 487 

this movement noted that the sells never moved directly outwards 
into the liquid but always at its surfaces. In Harrison's earlier 
studies he had attempted to cultivate fragments of the neural tube of 
frog embryos in hanging drops of serum. No evident activity was 
observed. Success was attained only when these fragments were 
placed in lymph which clotted about them. In studying the move- 
ment of the cells in lymph Harrison then noted that the cells moved 
always in contact with the fibrin fibrils, and in his later analysis of 
the culture of the Lewises he found the cells migrating always in 
contact with the surface of the cover glass or on the free surface of 
the medium. Harrison^^ termed this phenomenon, as L. Loeb-'^ had 
done, "stereotropism." He considered these cells apparently highly 
complex systems whose mechanism for movement is regulated 
through such contacts. Both authors thought this a common prop- 
erty of all cells. 

After a careful study of the movement of these embryonic cells in 
liquid medium. I noted, however, that these cultures of the Lewises 
were applicable only for the movement and growth of the cells of 
the younger embryos. No activity is seen about the normal frag- 
ments of heart muscle or, other mesenchyme cells of older embryos 
at the surface of the liquid or in contact with any solid. As I 
have stated above, blood plasma or fibrinogen is the one common 
substance of the body capable of stimulating activity in them. 

From these facts it seems evident, therefore, that it is not solids 
in a general sense that are necessary for the movement of these 
cells, but specific adsorbing substances. The younger embryonic 
cells differ from the older ones in that they may move at the sur- 
face of the salt solution and liquid media as well as in the clot. 
The older cells had lost this property. This leads me then to 
analyze more carefully this movement at the surface of the me- 
dium. I measured the position of the cells moving at the cover- 
glass surface with the micrometer of the microscope. I found that 
these cells were not in contact with the cover glass, as Harrison, 
Lewis-s and L. Loeb had thought, but that they lay often a con- 
siderable and measurable distance below it. Again, previous to the 
movement of the cells on the surfaces of the hanging drop, these 
surfaces change. They became covered by a film or scum, which 
made them appear leathery. The cells in every case moved, grew 
and divided in this film of material. This substance is not the 
coagulating substance "L" liberated by the other cells. It is son e- 
thing new, which acts evidently in place of and antagonistically to 



488 The University Science Bulletin. 

the fibrinogen. This substance formed early in the life of the cul- 
ture. It spreads rapidly to cover the whole of the medium. The 
cells later invade it as they invade the plasma. It differs from the 
fibrinogen in that it is a much more active stimulant. The cells 
form no insoluble compound with it. They grow actively for a time 
within it and invariably disintegrate. In the plasma cultures of 
these younger cells, a part of the cells may at first invade the clot. 
They soon leave the fibrinogen, however, for the surface film which 
has stronger affinities for them. Single cells may also liberate this 
substance. 

While I do not know the exact nature of this substance, it is 
interesting to note here that this substance disappears from or its 
action is inhibited about fragments of undifferentiated mesenchyme 
and the heart-muscle tissue in the cultures between the ninth and 
fourteenth day of the incubator life of the chick embryo. This is 
not true, however, of other tissue. I have identified it in the epider- 
mis and the liver of these embryos as late as the eighteenth day. 
Felix has shown the new tubules developed in the kidney of man as 
late as ten days after birth. While these studies are yet incom- 
plete, they do indicate that the disappearance or the inhibition of 
the action of this substance is not regular, but takes place at irregu- 
lar times in different tissue of these higher animals, and that this 
disappearance corresponds to the forms taken by the developing 
embryo and adult. 

What might be the exact chemistry of this substance remains to 
be seen. It seems most plausible, however, that the secrets of the 
building of the body will be found in the physical properties of it. 
These physical properties are, first, a strong affinity for water, 
and, second, an ability to stimulate an excessive metabolism. 

From the above observation it became evident, therefore, that the 
cells of the body are not in a continuous state of activity. The 
activity of early life is dependent completely upon the synthesis of 
a substance which removes the "L" substance froiH the cells. This 
substance is gradually superseded in later life by fibrinogen. Fibrin- 
ogen combines with "L" to form fibrin, an insoluble compound. In 
the later periods life centers about the functioning or differentiated 
systems. To prove this fact more definitely it became of interest to 
study the effect of a flowing stream of serum upon rhythmically con- 
tracting cells and the dedifferentiated muscle cells and the cells of 
connective tissue. 



Burrows: Study of Body Cells. 489 

THE CONTRACTING HEART MUSCLE CELLS. 

The isolated contracting heart-muscle cells which occasionally 
develop in these cultures from cells migrating from the fragments 
contract with a rhythm like that of the whole hearts or fragments 
of the heart transferred to the cultures. Since they occur most 
infrequently, contracting fragments have been used chiefly for this 
study. 

The rhythm of the fragments and the whole hearts of younger 
embryos may never cease when transferred from the chick embryo 
to the medium. If it does it commences at once again as soon as 
the temperature for it is restored. During the first few minutes 
or hours this rhythm is regular; and in the case of the whole hearts 
and fragments of the ventricle, it is the same as that which occurs 
normally in the body. After a short time, however, it becomes 
irregular. There are periods of activity followed by periods of 
complete rest. These periods of activity, as I have previously de- 
scribed them, are ushered in by rapid, strong contractions. These 
contractions gradually decrease in amplitude and rate to the period 
of complete rest. After a short rest period the active rhythm again 
intervenes, and so on; their irregular rhythm may continue for as 
long as eight or nine days in a single hanging drop. 

This slowing of the rhythm and rest, I concluded, was due to the 
temporary accumulation of waste products, the temporary lack of 
nutrient substances, or both. After these waste products had slowly 
diffused away and nutrient substances had moved in, the heart 
became active again. To prove this I placed several such fragments 
in a specially devised culture chamber. This chamber was arranged 
so that serum could be made to flow continuously along a cotton 
wick, the fibers of which transversed the layer of plasmatic medium. 
By this means it was possible to continuously wash the medium 
about the fragment. This culture has been called the ''wick cul- 
ture."-^ In such cultures the rhythm of the fragments remains 
regular during the time the serum is flowing, and this regular rhythm 
continues often for many days or until the protoplasm of the cells 
is otherwise destroyed by infection, etc. A careful comparison of 
the irregularities in the case of the contracting heart-muscle cells 
with those of the migrating ones shows interesting differences. In 
the case of the migrating and growing cells there are no intermittent 
rest periods. Migration commences after a latent period. It con- 
tinues actively for a time; then gradually ceases. In the same cul- 
ture there is no second recovery period. 



490 The University Science Bulletin. 

Again it is interesting to note that the changes in the clot are 
different in the case of these two activities. The migrating cells 
liberate a substance which occasions a coagulation of the clot. 
About the contracting fragments from which no cells are growing 
no such changes are seen in the clot. Contraction in the isolated 
cells commences after they have ceased to migrate and show these 
changes. 

The substance or substances liberated by the heart which pre- 
vents its contraction and which are evidently concerned with this 
act of contraction are soluble substances. They can be washed 
away with serum. The evidence gleaned from the above studies 
on the migrating cells indicates that the substance or substances 
which occasioned their migration are insoluble in the medium. They 
are adsorbed or chemically combined with the fibrinogen to form 
fibrin. The substances which accumulated to stop the activity of 
the heart is not the "L" substance noted above. 

For the testing of the connective tissue and undifferentiated heart- 
muscle cells I have used a glass culture chamber (plate LII) instead 
of the one described in the previous paper.-'' Otherwise the technic 
was the same. 

During the course of the study of the contracting heart-muscle 
■cells in 1912 I had thought that the migrating cells move more rapidly 
.against the stream and that they became more dispersed in these 
cultures than in the simple hanging drops of medium. These differ- 
-ences were observed, however, in but a few cultures. Later I noticed 
that the thickness of the layers of medium also affected the move- 
ment of the cells. The cells migrating from 1 mm. thick fragments, 
which had been placed near the edge of the hanging drop, moved 
more actively in the thin edge'^ than in the thicker parts of the layer 
within.^ Regulating these conditions in the "wick cultures," I found 
that the flowing serum in no way affected the migrating heart- 
muscle cells, nor did it effect in, any way the movement of the 
epithelial and connective-tissue cells in general. The only cells 
:affected were the leucocytes and lymphocytes. In the simple hang- 
ing-drop cultures it takes several transplants to effect a complete re- 
moval of the leucocytes and lymphocytes from fragments of bone 
marrow, lymph gland and spleen. In these "wick cultures" I found 
fragments of spleen entirely stripped of these cells after seventy-two 
hours. The leucocytes and lymphocytes had accumulated in masses 
■£it tangled parts of the wick. 

The conditions which regulate the movement of these latter cells 



Burrows: Study of Body Cells. 491 

are evidently different from those which control the movements of 
the fixed tissue cells of the body. Migratory movements and growth 
of the heart-muscle cells are effected by a substance which is insol- 
uble in circulating body fluids. Growth in these cells is therefore a 
purely physico-chemical reaction which proceeds to a condition of 
static equilibrium. For growth to take place it is necessary that 
this substance or these substances be removed. As they increase all 
activity ceases. This substance or substances I have designated 
as "L." 

DISCUSSION AND CONCLUSIONS. 

From these observations there seemed little doubt, therefore, that 
the dynamic state of the organism is not in any sense an indication 
of a similar dynamic state in the cell. It is a product of the organi- 
zation of the body. In early life the dynamic state is associated 
with the formation of a substance or substances which combine with 
or otherwise remove the "L" substance of the cell and occasion an 
active metabolism within them. This substance has strong affinities 
for water. It is thus directly concerned with the early building of 
the organism. It stimulates not only an excessive metabolism in 
these cells, but also may occasion the primary swelling of the mass. 
This substance disappears, or rather it ceases to be recognizable, in 
the early period of development. Subsequent to this period, life be- 
comes subservient to new organizations. The new organizations, the 
general nature of which I have illustrated in the study of the rhyth- 
mical contractions of muscle cells, are not unique, however, for the 
tissue. They are peculiar to all functioning cells. The nerve fibers 
are stretched between the brain and an end organ. Adrian has 
shown that the "all or nothing" law holds for this tissue like it does 
for the heart. The same is true for the glands. The secretory cells 
are cells which have a free end and one attached to a basement 
membrane. Stop up the ducts of one of these glands and these cells 
undergo atrophy. Such observations are wholly in line with the gen- 
eral facts which are Imown concerning development. In man the 
kidney may form tubules up to the tenth day after birth. Other 
organs cease their progressive growth much earlier. No careful 
studies are known concerning the time the heart cells cease to divide. 
In the later embryonic period, at least, all growth in the heart is 
represented by an increase in the size of the fibers rather than an in- 
crease in the cells. In the kidneys and glands it manifests itself in 
a dilation and increase in length of tubules and a flattening of their 



492 The University Science Bulletin. 

lining cells. In the wound the exudate is the active stimulus for 
growth. It stimulates metabolism through a direct affinity of the 
"L" substance for the fibrinogen. "L" combines with fibrinogen to 
form fibrin. The fibrin then becomes the extracellular or collagen 
fibrils. The cells remain active and move towards the center of the 
exudate until the reaction is satisfied. The end of activity is the 
completion of the reaction. It is the scar for the connective tissue 
and continuity, and the limiting membrane for the epithelial cells. 
Through the analysis of the simple act of changes of shape and 
the movement of the connective-tissue cells it has been possible to 
show that these acts are not the result of any complex cellular or- 
ganization, but a simple reaction between substances in the environ- 
ment and a substance of the cell. The movement is the result of the 
cohesion of this substance for the cell on one side and specific sub- 
stances in the medium on the other. In this act of locomotion the 
cell supplies the energy; the mechanism otherwise resides wholly in 
the environment. Again, in other studies I have been able to show 
that the syntheses for growth are not a part of these energy-pro- 
ducing reactions, but they are separate reactions. The connective- 
tissue cells in the adult organism are widely separated in a mass of 
fibrillar substance. These cells migrate into the plasma, but they 
do not grow. In the plasma culture, growth is peculiar alone to the 
more cellular fragments of embryonic tissue, granulation tissue or 
sarcomata. By, the use of embryonic extracts it is possible to stim- 
ulate the metabolism of these connective-tissue cells. Under these 
conditions the widely separated cells of the adult fragments will 
grow and divide. About the more densely cellular fragments such 
stimulation is not necessary. It is harmful. It leads to the destruc- 
tion of the cell, a breaking down of the proteins of the protoplasm. 
If these same fragments are teased apart so the cells become more 
dispersed, growth ceases again. For growth to take place it is neces- 
sary that the cells be either crowded or excessively stimulated. The 
important factor for this reaction is evidently the concentration of 
certain products of their metabolism. The concentrating of such 
products may be induced by the crowding of cells or increasing the 
rate of their production. These cells liberate not only the "L" sub- 
stance, but also CO2 and H2O. How many other substances are 
formed when the reaction is proceeding has not been determined. 
Protein synthesis is not a part of the ordinary metabolic reaction of 
the cell. It is secondary reaction depending upon and obtaining the 



Burrows: Study of Body Cells. 493 

energy necessary for it from the energy-producing reaction of the 
cell. The extracellular fibers of the connective tissue are not secre- 
tions of the cells any more than bone or cartilage are of this origin.^ 
These fibers are the combination of proteins formed elsewhere in the 
body and the ''L" substance of these cells. 

There is no reason to believe, therefore, that protein synthesis is 
a part of the metabolism of the cell. It is something different. It 
is a form of work produced. Upon it growth depends. Growth has 
never been shown to be of a simple chemical or physical nature. 
It is the result of the careful utilization of energy. It takes place 
against the forces of nature. Protein synthesis, like muscular con- 
traction, is only a form of work peculiar to body organizations and 
not cellular organizations. 

In 1917,^*^ and again in recent experiments,^ I have shown that 
the development of ryhthmical contraction is not associated with 
any fundamental change in the cell. This is a property peculiar to 
any of the mesenchyme cells of early embryonic life. It occurs in 
the fragments of this tissue and in the cells which migrate from 
them. Its development is the result of a chance relationship of the 
cells to medium. This relation is wholly dependent upon the 
physical peculiarities of the coagulation of the plasma clot. The 
isolated cells which develop rhythmical contraction are those cells 
which become stretched through a serum cavity between the sur- 
face of the medium or a cellular tissue fragment and the end of 
fully formed fibrin fibrils. No contractions develop until these 
fibrin fibrils are fully formed. What is true for the isolated cell 
is also true for the fragments. Fragments of the heart of young 
embryos contract at once when removed to warm medium of 
the culture. Those from older embryos fail to show this change. 
Rhythm develops in these latter fragments only after the border 
cells have moved out or the process of coagulation is completed. 
The end of the cell in contact with the fibrin is in metabolic equi- 
librium. These cells imbedded in fully formed clot cease all ac- 
tivity. The cells floating in serum also show no change. The only 
active part of these cells differentiated for contraction is the free 
end in contact with the cellular fragment or the surface of the 
medium. This end suffers a decrease in surface tension. Such a 
decrease in surface tension is associated with electrical changes at 
this end of the cell and a stretching of the cell. If such changes 
continue, one of three conditions must result: the cell will be torn 
loose or in two, or there will be an explosive breakdown of this 



494 The University Science Bulletin. 

surface-tension-lowering substance. That such an explosive break- 
down is peculiar to the contraction is clearly indicated by corre- 
lating these observations with those of Fletcher. Fletcher has found 
that lactic acid is liberated during the contraction phase of muscle. 
I find that lactic acid increases the surface tension of the cells. It 
causes the cell to contract. This lactic acid disappears again dur- 
ing the intermittent rest period. The cell again returns to its former 
state. The process is rhythmical. The process, as it is evident, is 
somewhat similar, therefore, to the phenomenon described by Bredig. 
Bredig showed that when a ten per cent solution of H2O2 is placed 
over the surface of pure mercury, a film of mercury peroxide forms 
at the surface. This leads to electrical changes in this system. 
Under appropriate conditions the peroxidate breaks down again 
to mercury and oxygen. Then the layer of peroxidate reforms. The 
process is repeatable or rhythmical. Bredig and his students further 
find that many conditions which alter rhythmical muscular con- 
traction also alters the activity of this model.^^ 

Rhythmical muscular contraction is not, therefore, as Bayliss sug- 
gests, the result of rhythmical stimuli, but evidently the result of 
an explosive breakdown leading to alternate changes, not only in 
surface tension, but in the electrical conditions at opposite ends of 
the cell. Bernstein'^" several years ago had given definite evidence 
to show that the energy of muscular contraction is surface energy. 
He found that muscles suffer the thermal changes peculiar alone to 
surface energy. 

In such organization, as it -is well recognized, there is no reason 
that such an explosive breakdown should always occur. Such is 
possible only when the decrease in surface tension at the free end 
or the electrical changes are of such a degree to allow a current to 
pass through the resistent cell. This change at the free end may 
cease before such is possible. For a breakdown to take place other 
special conditions or stimuli must be present. That this is the con- 
dition of most of the functioning tissue of the body has been well 
proven by the studies of tissue autonomy. In man, whether any 
muscular tissue, other than the nodes of the auricle, are autonomous, 
like the cell of the culture, I think is questionable. At the same 
time it is evident that any functioning tissue may develop such a 
condition. Whether this is what has happened in many of the 
nervous affections might be a problem worth investigating. It is 
not surprising, however, that in other animals the autonomy is 
centered in other tissues. Carlson^^ finds that the rhythm of 
the heart of the limulus is not automatic, but centered in certain 



Burrows: Study of Body Cells. 495 

ganglion cells. In men I think it is very questionable whether the 
nervous system, any more than the striated muscle or gland cells, 
respond without the aid of external conditions. It is upon this fact 
that coordination depends. 

There is no evidence from the above observations that the cells 
themselves undergo any absolute changes during these fundamental 
changes in the body. For a good many years it has been well 
known that bone and cartilage are products of a given organiza- 
tion of the part and not a product of the cell. It has been shown, 
for instance, that bone will develop in the pelvis of the kidney if 
the blood vessels to that organ are ligated. Asami and W. DocTi^^ 
repeated these experiments in the laboratory and proved completely 
the existence of these changes, as the other authors had shown. In 
the tissue culture the cells migrate readily, not only from fragments 
of interstitial tissue, tendons and fascia, but also from fragments 
of bone and cartilage. The bone cells and cartilage cells behave in 
every way like the other connective-tissue cells. The bone and 
cartilage remains behind like the extracellular fibrils. In the same 
way the adult muscle fiber will not react in the plasma, but I have 
seen the nucleus and sarcoplasm migrate out and leave the adult 
fiber behind. In the plasma this mass forms perfect connective- 
tissue cells. These cells react also in every way like the ordinary 
connective-tissue cells. 

Such simple fluid systems may not only suffer changes in shape 
and differential changes of tension at various points on their sur- 
face, but thej^ may also suffer additions and probably subtractions 
from themselves. In the body differentiation is not only the result 
of mechanical changes initiated and controlled by the environment, 
but it may be also chemical in nature. As the above studies of the 
skin epithelium show, these cells do not suffer fundamentally from 
the connective-tissue cells. For metabolism to take place within 
them it is necessary that the ''L" substance be removed from them. 
This "L" substance is not different in these cells from that of the 
connective-tissue cells. These cells differ from the connective tissue 
in that they also contain a proteolytic ferment. The pancreas cells 
again differ in that they also contain a fat-splitting ferment. They 
often fail to migrate into the clot. They lead rather to the rapid 
splitting of the fat of the plasma to fatty acid crystals. This pre- 
vents also the migration of the connective-tissue cells from these 
fragments into the medium. So in each case the various tissues 
differ as they contain their own peculiar added products. The fun- 



496 The University Science Bulletin. 

damental reaction of the fixed tissues otherwise remains the same. 
The only exception to this rule is to be found in the wandering cells. 
These cells have not been observed to grow in the cultures. They 
have also lost their ability to cause a true coagulation of the plasma. 
They occasion the gelation of the clot and can move only in contact 
with this jelly like mass, but they cause no fibrin formation. They 
owe their spherical shape to their inability to form true surfaces in 
the medium. They move by a mechanism different from the fixed 
tissue. They can invade these fixed tissues in the presence, at least, 
of an exudate. When this disappears they tend to move back into 
the lymphatics and blood capillaries. They do not repel strongly 
the moving fixed tissues like the fixed tissues repel each other. They 
have lost the property to form the "L" substances. I say they have 
lost it because it is present in the mother cells from which the wan- 
dering cells arise. 

To what extent this chemical differentiation is reversible like that 
peculiar to -the mechanical form I have not definitely determined. 
There is evidence to show that the mesenchyme cells may arise from 
epithelial cells even in late embryonic life, but for the most part 
these epithelial cells maintain their chemical peculiarities for a 
long time in cancerous growths and in the cultures. I have seen 
heart-muscle cells assume the characteristics of large mononuclear 
cells. The reverse has not, however, so far been proven. Again 1 
have seen liver cells, after repeated transplantation, behave in part 
at least like the connective-tissue cells. There is no reason to be- 
lieve, however, that such chemical dedifferentiation may not occur 
and maintain in the proper environment. 

The cells of the organism are not, therefore, highly complex sys- 
tems. They are not equipped to lead an independent existence. 
They do not age. They have no organization for work. They pro- 
duce the energy, but the work or their various manifestations of life 
is dependent wholly upon external conditions about them. The 
only form of work depending on an evident internal organization is 
cell division. The forces active in the process center about the 
centrosomes. There is no evidence, however, that activation of these 
centers, or even their formation, is controlled from within. The 
centrosomes develop probably alone in response to external stimuli 
(see Hertwig, Meade, Morgan, J. Loeb, E. B. Wilson^ and others). 

Energy production in these cells is again wholly dependent upon 
organization, or the presence of specific substances which split or 
otherwise make the removal of the "L" substances possible. Their 



Burrows: Study of Body Cells. 497 

activity in early life is dependent upon the synthesis of a substance 
which combines or otherwise removes the "L" substance. In later 
life it is dependent upon a mechanical differentiation. This form 
of synthesis in later embryonic life forms a substance which is 
fibrinogen or closely akin it. This forms an insoluble compound 
with "L" rather than the soaplike substance of the earlier period. 
In the proper environment it occasions, through its coagulation, the 
form necessary for the dynamic state. 

By these observations it has been possible, therefore, for the first 
time to define differentiation in other than morphological terms, 
and to compare function with growth. The growth of the undiffer- 
entiated cells of early life is the result of a special synthesis. The 
development of function is the result of a slightly different one. 
Differentiation is quite different, therefore, from what it has been 
conceived to be. 

While these observations reduce growth and function to simple 
physicochemical formulation, they give no hint as to the cause for 
this change in synthesis from early to late life. Any substance 
which removes the "L" causes an immediate loss of mechanical 
differentiation. The heart-muscle cells in contact with the fibrin 
become simple mesenchyme cells. Only under proper mechanical 
conditions can they redifferentiate. What is true for the heart is 
true for the glands. In the cultures the gland cells stretch to form 
membrane like the skin. They lose the form necessary for function. 

These facts, again, do not explain chemical differentiation. In 
the above observation I have also not discussed all the deficiency of 
these cells. Besides the lack of any organization for work and any 
means within themselves to allow the energy-producing reaction to 
proceed under the ordinary conditions in nature, they are also bereft 
of the property of using the crude material of nature for this energy- 
producing reaction. While energy production in the body is derived 
chiefly from H and C, these cells cannot use the sugar carried to 
them by the blood stream without intervention of substances from 
the pancreas. The organization of the whole or certain of its parts 
are again essential for another of their important needs. 

It is upon this last deficiency that the chemical syntheses peculiar 
to differentiation must depend. The body is a machine operated like 
any other machine in nature. Its metabolism has no complexities, 
as most biologists would have it. It is a machine which is able to 
produce energy and transform it into work. It has been possible to 
locate these different reactions. The energy is produced in the cell; 



498 The University Science Bulletin. 

its transformation is under the control of the environment. Protein 
synthesis is in no way to be confused with the energy-producing re- 
action. In the body, anything which increases metabolism will lead 
to growth. Increase this still more and protein destruction results. 
This accounts for the appearance of split products of protein in 
muscle which has suffered excessive stimulation (see Bayliss for lit- 
erature) and the destruction of the tissue by strong growth stimuli 
such as X-ray, radium, coal tar, arsenic, etc. 

Such a machine can build itself in the manner that it does because 
it utilizes the products of its energy-producing reaction for its build- 
ing. The building is only the sequence of change its original funda- 
mental structure undergoes to produce the final necessary work — 
the work of supplying an adequate amount of fuel for a certain pe- 
riod of life and supplying the egg with an adequate supply of this 
material (yolk) to carry it through the early period of the develop- 
ment of the whole. This yolk supply decreases progressively. This 
decrease leads to the changing character of the syntheses. 

The extracellular deposits which I have described above as im- 
portant for the organization peculiar to the dynamic state of later 
life are evidently the substratum recognized by Child. They are the 
result of protein syntheses peculiar to the organization of a certain 
period of the development of the whole. They must be the result 
not only of mechanical but also chemical changes in the environ- 
ment. The picture of the organization as it is seen through the study 
of the cells is in no sense the picture of one continuous metabolic 
change. It is the primary building of a heterogenous system fol- 
lowed by the gradual decline of this system to a state of static 
equilibrium. Elemental life and elemental death are not compar- 
able to systemic life and systemic death. The body is the necessary 
cycle that these cells may preserve their kind. For the active growth 
of the early period of the development the cell draws upon the yolk 
or the mother for its supply of those substances necessary for this 
growth. The disappearance for this supply of material is the ap- 
pearance of the second, or the functioning, period. The building of 
this period is completed within ten days after birth in man, except 
for the laying down of the nerve sheath. The syntheses peculiar for 
this building are the result of the disappearance of the substances 
carried in the yolk. The important factors for differentiation are 
not to be found, therefore, in the primary reactions of the cells, but 
in the deficiencies of these cells. The important deficiency which 



Burrows: Study of Body Cells. 499 

has to do with differentiation is their inability to utilize the crude 
food materials supplied by nature. This deduction we are forced 
to believe, however, not only from the above observations, but from 
a host of already carefully accumulated facts. If the first two 
blastomeres of many lower forms are separated, two animals de- 
velop. The yolk is also separated by this process. Each animal is 
but one-half the size. Gudernatsch has shown that tadpoles fed 
upon thyroid differentiate within a few days, while thymus feeding 
delays this process at least for a very long time. Very large tad- 
poles may be thus developed. The ultimate heterogenicity thus de- 
veloped is essential for subsequent life and for reproduction. The 
egg cell builds a new system like the old (heredity), in that it has 
acquired from the old a supply of those substances necessary to 
carry it through the building of the new system, which is again ca- 
pable of preparing these substances from the crude materials avail- 
able in nature wdthout. The building results from the changing 
syntheses which result from the gradual decreasing yolk supply. 
The form of the building is dependent, therefore, primarily on the 
original constitution of the yolk. This is again dependent abso- 
lutely on the nature of the machine which produces it. Each ani- 
mal must, therefore, in each case reproduce its kind. The problem 
of heredity is thus reduced to pure physics and chemistry. 

Death in such a system may result, therefore, from the destruc- 
tion of essential parts, or the inevitable equilibrium of those forces 
which maintain the heterogenicity. This does not mean a funda- 
mental change in the cell. The cell succumbs as the result of this 
breakdown. There is no reason why any of these cells may not 
grow actively again if the organization is changed about it. Differ- 
entiation is both chemical and mechanical in nature. Chemical dif- 
ferentiation does not effect the fundamental energy-producing reac- 
tion of the fixed tissue cells. It is an indication of the chemical 
heterogenicity of the mature organism. The mechanical differen- 
tiation essential for the dynamic state is reversible. According to 
the above formulation, mechanical differentiation in early life is the 
result of the removal and decrease in certain materials of the yolk. 
If a change in organization suitable for active growth should take 
place in the organism, no such differentiation should follow. The 
cells suffering such changes must continue to grow as long as the 
body supplies the necessary substances. The body must supply 
those substances necessai-y for active growth as long as it survives. 



500 The University Science Bulletin. 

A rapid utilization of these substances must lead to a rapid atrophy 
and death, according, also, to the formulation given above. 

Such a reorganization, it is evident, cannot take place in early life 
except under the influence of most powerful external stimuli or de- 
velopmental defects. It becomes, however, more possible in later 
life when the normal forces which maintain the normal heterogenic- 
ity are waning. Cancer represents such a form of active growth. 
It is a disease peculiar to later life. It occurs earlier in connective- 
tissue areas than in epithelial tissue. The connective-tissue cells, 
according to the above observations, lose their property for inde- 
pendent growth earlier than the epithelial tissue. 

In previous publications ^ I have noted that cancerous tissue 
grows like that of younger embryos and liberates the same or a 
similar growth stimulus.^^ In the body cancer may result from 
congenital abnormalities, such as pigmented moles. It occurs more 
frequently in certain families of mice (Maude Slye). It follows the 
continuous application of many growth stimuli or substances capa- 
ble of effecting such a reorganization. In man, as pathologists agree, 
it follows most frequently upon long-standing chronic inflamma- 
tion (Billroth).=^« 

As has long been fully appreciated, the impediment which has 
stood in the way of advance in cancer has been the inadequacy of 
our knowledge of the cell and its relation to the whole. It has been 
the endeavor of the cancer laboratory in St. Louis to attack the 
problem from this routg. Cancer is not a parasitic disease. It is a 
disease which follows after long, continuous stimulation. Before 
cancer can be understood it is necessary that irritability and stimu- 
lation be reduced to simple terms. The essential conditions for 
mechanical dedifferentiation, or loss of function, and the production 
of an active growth of cells like that of cancer is the presence of 
substance capable of removing the "L." This "L" has many of the 
properties of the phospholipins isolated by Mills. These phospholi- 
pins are soluble in many lipoid solvents, and especially products of 
coal tar. We have studied the action of coal tar, and find that it be- 
haves in the tissue like the substance liberated by the cancerous 
tissue and the tissue of young embryo which is able to combine with 
or otherwise remove the "L" from the cells. Coal tar thus attracts 
the cells to it and effects their dedifferentiation. At first it occasions 
their disintegration. Later it becomes a less active solvent and 
occasions an active proliferation of these cells. It is capable not 
only of producing many of the symptoms of cancer by itself, but of 



Burrows: iStidv of Body Cells. 601 

effecting the ivurganization lu'cessary tor true cancer to develop. 
Long use of coal tar leads to tiie de\'el()pment of cancer.^'^ 

What I have hoped to present in this paper is not only the general 
picture of the cell the tissue culture has so far revealed, but methods 
by which the further details of these general problems may be at- 
tacked. From the above observations it is evident that the problems 
relative to life are not to be solved by chemical and morphological 
methods, but by the application of mechanism. Heredity, as we 
have seen, is dependent wholly on the chemistry of the system, but 
the importance of these chemical substances will not be understood 
until the mechanisms peculiar for the various manifestations of life 
have been isolated, their parts and the energy relation between these 
parts have been fully determined. Upon this latter knowledge rests 
also the ultimate control of cancer. Cancer, as it has been seen, does 
not arise from embryonal cells. It is not the mere displacement of 
cell (Ribbert) , but a specific reorganization of parts. Such a re- 
organization means a breaking down of the normal relation of cells 
in the adult. The normal heterogenicity is maintained through the 
fact that each kind of cell moves through the liberation of the same 
surface-tension-lowering substance. Each tissue repels its neighbor 
like each cell repels each other cell of the same or of another kind. 
It is a breaking down of this barrier through the synthesis of a new 
substance which conditions new surface relations. It is an invasion 
of connective tissue by the epithelium (Thiersch and Remak) . This 
synthesis becomes possible through the specific rearrangement of 
cells. Such rearrangements may result from developmental defects 
or through the action of stimulating substances. In the body not 
only quality but quantit}^ is an absolute factor. Systemic life and 
elemental life are not comparable. The cells do not age. Differen- 
tiation is not an age phenomenon. The law of the conservation of 
energy holds for the body like it holds for all natural phenomena. 
Advancement in biology is to be made through the development of 
methods for the study of the mechanics of these systems. Mor- 
phology and chemistry must be supplemented by these methods. All 
parts of these machines are not visible. Rhythmical heart-muscle 
contraction is not a specific chemical reaction. It is peculiar to a 
large number of animals. These animals differ chemically from 
each other. A watch is a watch whether it is made of gold or silver. 

The heart contracts as the result of a special arrangement of its 
parts and the specific dynamic properties of these parts, and not 
through the specific chemical constitution of the particular protein 
which compose its parts. 



502 The University Science Bulletin. 

BIBLIOGRAPHY. 

1. Child, Charles Manning. 1915. Senescence and Rejuvenescence. The 
University of Chicago Press, Chicago. 

2. Thompson, D'Arcy W. 1917. Growth and Form. University Press, Cam- 
bridge, London. 

3. Wilson, E. B. 1906. The Cell in Development and Inheritance. Second 
edition. London and New York. 

4 Huxley, T. H. 1863. Review of the Cell Theoiy. British and Foreign 
Med. Cir. Rev., vol. XII. 

5. Wells, H. Gideon. 1914. Chemical Pathology. Second addition. Phila- 
delphia and London. 

6. Bayliss, W. M. 1915. Principles of General Physiology. London. 

7. Burrows, M. T. 1913. The Tissue Culture as a Physiological Method. 
Trans, of 'the Congress of Am. Phy. and Surg., vol. IX, pp. 77-90. 

8. Burrows, M. T. 1913. Tissue Culture in vitro. XVII the International 
Congress of Medicine, General Pathology and Pathological Anatomy, Lon- 
don, pp. 217-237. 

9. Burrows, M. T. Titles Read May, 1923, to Appear in Oct., 1923, Proc. of 
the Society of Exp. Biology and Medicine, vol. XXI. 

10. Burrows, M. T. 1911. The Growth of Tissue of the Chick Embryo Out- 
side the Animal Body with Special Reference to the Nervous System. 
Journal of Exp. Zool., vol. 10, pp. 63-83. 

11 Burrows M. T. 1916-17. Some Factors Regulating Growth. Anat. 
Record, vol. 11, pp. 335-339. 

12. Fletcher, W. M. 1902. Tlie Relation of Oxygen to the Survival Metab- 
olism of Muscle. Jour, of Physiol., vol. 28, pp. 474-498. 

13. LoEB, Jaques. 1913. Artificial Parthenogenesis and Fertilization. Chicago 
University Press, Chicago. 

14. Morgan, T. H. 1901. Regeneration. New York and London. 

15. MoRG.\N, T. H. 1906. The Physiology of Regeneration. Jour, of Exp. 
Zool., vol. 3, pp. 457-500. 

16. WooLBRiDGE, T. C. 1893. On the Chemistry of the Blood, and Other 
Scientific Papers. The Crotian Lecture on the Coagulation of the Blood. 
London. 

17 Mills, C. A. 1921. Chemical Nature of Tissue Coagulins. Jour, of Biol. 
Chem., vol. XLVI, pp. 135-165. 

18. Hertzler, a. E. 1904. Peritoneal Adhesions, Their Cause and Prevention. 
Tr. West. Surg. Assoc, vol. XIV, p. 76; Anat. Rec. 1915, vol. IX, p. 83. 

19. Hertzler, Arthur E. 1919. The Peritoneum. Vol. 1, chap. V. St. Louis. 
20 Baitsell G. a. 1916. The Origin and Structure of Fibrous Tissue Formed 

in Wound Healing. Jour. Exp. Med., vol. XXVIII, pp. 739-756. 

21. Burrows, M. T. 1915. An Attempted Analysis of Growth. Anat. Record, 
vol. 9, No. 11. And, The Tissue Culture in Cancer. Proc. Second Pan- 
Amer. Sci. Congress, Washington, section VIII, part 2, pp. 494-496. 

22. Burrows, M. T. 1912. Rythmische Kontraktionen der isolierten Herz- 
muskelzelle ausserhalb des organismus Mumchener medizinischen. Wo- 
chenschrift, No. 27, pp. 1-10; and Science, N. S., XXXVI, pp. 90-92. 

23. Burrows, M. T. 1913. Wound Heahng in vitro. Proc. of the N. Y. Path. 
Soc, N. S., vol. XIII, Nos. 5 and 6. 

24. M. R. and W. H. Lewis. 1911. The Cultivation of Tissues from Chick 
Embrvo in Solution of NaCl, CaCl2, KCl and NaHCOs- Anat. Rec, vol. 
5, pp. 277-294. 

25. Harrison, R. G. 1911. On the Stereotropism of Embryonic Cells. Science, 
N. S., vol. XXXIV, pp. 279-281. 



Burrows: Study of Body Cells. 503 

26. Harrison, R. G. 1914. The Reaction of Embryonic Cells of Solid Struc- 
tures. Jour, of Exp. Zool., vol. 17, pp. 521-544. 

27. LoEB, Leo. 1922. Agglutination and Tissue Formation Science N. S., 
vol. LVI, pp. 237-240. 

28. Lewis. W. H. 1922. Is Mesenchyme a Syncytium? Anat. Record, vol. 23, 
pp. 177-184. 

29. Burrows, M. T. 1912. A Method of Furnishing a Continuous Supply of 
New Medium to a Tissue Culture in vitro. Anat. Record, vol. 6, pp. 
141-144. 

30. Burrows, M. T. 1917-'18. A Note on the Mechanism of Heart-muscle 
Contraction. Am. Jour. Physiology, vol. 45, pp. 556-557. 

31. Bredig, S., und WEiNMA-iTiR, J. 1903. R, periodische Kontaktkatalyse. 
Zeitsch fur Phys. Chem., B. 42, S. 601-611. And, Bredig, S. 1907. Bio- 
Chem. Zeitchr., B. 6, S. 283. 

32. Bernstein, J. 1908. Uber die Temperaturcoeffizienten der Muskelonergie. 
Pfluger's Arch., B. 122, S. 129-195. 

33. Carlson, A. J. 1904. The Ner\^ous Origin of the Heartbeat in Limulus 
and the Nervous Nature of Coordination or Conduction in the Heart. 
Amer. Jour. Phys., vol. 12, pp. 67-74. 

34. AsAMi, GoicHi, .\ND Dock, William. 1920. Experimental Studies on Hy- 
perpastic Bone Formation. Jour. Exp. Med., vol. XXXII, pp. 745-766. 

35. Burrows, M. T. 1923. The Experimental Production of Malignant Ulcers 
in the Rat. Mo. State Med. Jour., vol. XX, pp. 145-147. 

36. EwiNG, J. 1919. Neoplastic Diseases. Philadelphia and London. 

37. L. H. Jorstad. A Study of the Behavior of Coal Tar in the Tissues. Proc. 
of the Soc. Exp. Biol, and Med., Oct. 1923. 



504 



The University Science Bulletin. 



PLATE LII. 




The "wick culture" chamber. 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 20— October, 1922. 

(Whole Series, Vol. XXIV, No. 20.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

Notes on the Biology of Curicta (Heteroptera), 

Grace Olive Wiley. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE. KAN. 



Entered at the post office in Lawrence as second -clas.s matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XIV. I October, 1922. [No. 20. 



Some Notes on the Biology of Curicta* from Texas. 

BY GRACE OLIVE WILEY. 

BEHAVIOR OF ADULTS AND NYMPHS OF CURICTA. 

I COLLECTED several pairs of adult Curicta and placed them all 
in one glass to take home alive. When I reached home most of 
them were mating. These were removed and placed in separate 
glasses, and remained paired for several hours. 

In mating, the male takes a position to one side of the female, and 
usually to the right. If to the right, he hooks his left anterior tarsus 
over her head; if to the left, the right fore tarsus is used. 

Both nymphs and adults seem fond of getting out of the water 
and lying close to the ground, where they are hardly discernible. I 
have found adults almost a foot from the water's edge, in tangled 
plant roots and under rotten pieces of wood. Search for eggs laid 
in nature provided fruitless. One pair was mating. I am half in- 
clined to believe the eggs are laid in soft mud.f 

The nymphs are very agreeable, in that they do not feed upon 
others of their kind, even when hungry. They like small notonec- 
tids, corixids, small carabids, fresh-water shrimp, and such. They 
refused small minnows, however. It is not uncommon to see three 
feeding quietly on one shrimp, or two feeding on one small beetle. 
They are very fond of mosquito larvae. 

EGGS. 

Size. About 1.75 mm. long; width a trifle more than .75 mm.; diameter of 
crown a little less than % mm. Rosette of filaments at tip, numbering 15; 
length of filaments almost 1 mm. 

* Curicta drakei Hungerford. 

t Have now found the eggs deposited in the tissue of dead plant stems with only the- 
crown of filaments visible. 

(507) , .•^"^ '■ 



508 The University Science Bulletin. 

Shape. Elongate oval, one end slightly tapering and rounded, the other 
smaller, sloping somewhat obliquely and bearing a crown or rosette of fifteen 
long filaments. 

Color. Creamy white, with filaments somewhat yellowish, center of crown 
darker. 

Seven eggs were laid by one female twelve days after mating. 
These were not inserted in soft wood or in plant tissues, althougii 
both were available. The female was lying close to the wet sand 
and the eggs were laid on their sides on the sand without any regu- 
larity or order. The next day there were three more eggs, and in 
three more days I found six eggs hidden among the roots of water 
plants and in slimy accumulations in crotch of a dead twig. 

DESCRIPTION OF FIRST-INSTAR NYMPH. 

Form elongate oval; very much like adult, only broader in proportion to 
length. Head large, much narrower than prothorax and as wide as long, ex- 
cluding the rostrum. Eyes globular, small. 

Anterior femur quite robust and armed with a single median tooth, plainly 
\isible and closer to the base than the apex of the femur. 

Anterior coxae about half the length of their femora, and very robust. In- 
termediate and posterior legs short. One-segmented tarsi. 

Color when first hatched pale straw yellow, becoming darker with brownish- 
black markings. Number of days of first instar, twelve. Entire length of in- 
sect from tip of beak to end of respiratory tube, 5 mm.; width across abdo- 
men at widest part, 1.50 mm.; width across eyes, .75 mm.; width of shoulders 
at base of head, 1.15 mm.; length of prothorax on median line, 1 mm.; length 
of respiratory tube, a trifle more than .50 mm.; length of anterior femora, 1.40 
mm.; length of anterior tibiae, .70 mm.; length of anterior tarsi about .25 mm.; 
length of anterior coxae, .70 mm.; length of intermediate femora, 1.40 mm.; 
length of intermediate tibiae, 1.00 mm.; length of intermediate tarsi, .20 mm.; 
length of posterior femora, 1.25 mm.; length of posterior tibiae, almost 1.50 
nmi.; length of posterior tarsi, almost .50 mm. 

Since writing the above I have reared the insect through from the 
beginning. There are five instars. 



510 The University Science Bulletin. 



PLATE LIII. 

The genus Curicta was represented in the United States by Curicta howardi 
Mont., described from a single specimen taken at Victoria, Tex., years ago. 
Nothing was known concerning the biology of these bugs. The sketch of the 
egg and first-instar nymph are therefore of interest. 



Wiley: Biology of Curicta. 
PLATE LIII. 



511 




l^t. In*ta.T Cur (tta. . hoiwardr. 












K(^<^ **\ CuT»«i,\aL . 



i 



THE 

KANSAS UNIVERSITY 

Science Bulletin 



Vol. XIV, No. 21— October, 1922. 

(Whole Series, Vol. XXIV, No. 21.) 

ENTOMOLOGY NUMBER V. 



CONTENTS: 

Biology and Morphology of Lepyronia quadrangularis 

(Say) — Homoptera, Cercopid.e Kathleen Doering. 



PUBLISHED BY THE UNIVERSITY 
LAWRENCE, KAN. 



Entered at the post office in Lawrence as second-class matter. 

9-4522 



THE KANSAS UNIVERSITY 

SCIENCE BULLETIN 

Vol. XI V.J October, 1922. [No. 21. 



Biology and Morphology of Lepyronia quadrangularis 
(Say) — Homoptera, Cercopidse. 

BY KATHLEEN DOERING. 

Submitted to the Department of Entomology, University of Kansas, in partial fulfillment of 
the requirements for the degree of Master of Arts. 

INTRODUCTION. 

THE purpose of this paper is twofold; first to add to the limited 
data concerning the biology of the Cercopidse; and secondly, to 
contribute to the morphological knowledge of the family, and inci- 
dentally of the order. The subject of this paper was suggested by 
Dr. Paul B. Lawson. Since Lepyronia quadrangularis (Say) is one 
of our most common spittle bugs, it was thought that a study of its 
biolog\", habits and morphology would be well worth while. Later 
when a review of the literature revealed the meagerness of both bio- 
logical and morphological details concerning other species in the 
family, the necessity for this work was magnified. 

By means of the morphological studies the writer has attempted 
to accomplish two things: First, since little work has been done on 
the morphology of any cercopid, to give a detailed description of the 
external anatomy, merely for the morphological interest involved; 
and secondly, to contribute a small share, if possible, in determining 
the relationships of the families within the order. A thorough 
knowledge of the morphology of all families in a given order seems 
necessary to correctly determine their phylogenetic relationship. In 
the literature, however, they are usually determined on the basis of a 
comparison of certain special parts of the body of a single species 
from each family. But since the species, or at least the genera, may 
vary within a family, some having primitive head sclerites and spe- 
cialized genitalia, while others are just the opposite, it follows that 
a fairly thorough knowledge of the entire structure of the body 

(515) 



516 The University Science Bulletin. 

should be obtained before such relationships are determined. To 
this end this small contribution is made. 

The writer wishes to express her appreciation to all who have as- 
sisted in this work: To Prof. S. J. Hunter for his interest and readi- 
ness to help; to Dr. Paul B. Lawson, under whose direction the work 
was done, and who, at the sacrifice of his own research time, gave 
most helpful advice and criticism; to Dr. Grace M. Charles for the 
correct identification of most of the host plants ; to Lucy M. Hack- 
man, Dr. H. B. Hungerford and P. A. Readio for their kindly help 
and criticism, and to all others who in any way have assisted in this 
work. 

THE SYSTEMATIC POSITION. 

FAMILY CHARACTERISTICS. 

The Cercopidse are distinguished from the other families of Ho- 
moptera by having three-segmented tarsus, two ocelli, the antennse 
inserted in front of and between the eyes; the prothorax not pro- 
longed backward, and the hind tibiae armed with one or two stout 
teeth, with two rows of short, stout spines at the tip. 

SUBFAMILY CHARACTERISTICS. 

The Cercopidse are divided into two subfamilies — Cercopinae Am. 
& Serville, and Aphrophorinse Am. & Serv. Lepyronia quadrangu- 
laris (Say) comes within the latter. According to Ball (1898) the 
Aphrophorinse are separated from the Cercopinae by the following 
characteristics: Anterior margin of the pronotum angulate; head 
equaling or almost equaling the pronotum in width; elytra com- 
pressed behind, rarely reticulate. 

HISTORICAL REVIEW. 

This insect has been described under various genera. It was first 
made known to science by Say in 1825 as Cercopis quadrangularis. 
In 1831 is! was put in the genus Aphrophora by Say, and in 1851 in 
the genus Ptyelus by Walker. Amyot and Serville (1843) described 
it under the genus Lepyronia, which name it now holds. 

GENERIC DESCRIPTION. 

The original description of the genus by Amyot and Serville is 
as follows: "Corps court et remasse. Tete en cone arrendi anteri- 
eurement, sans carene longitudinale mediane sur le vertex ni sur le 
front. Elytres bombees, en ovale court et en forme de coquille. Les 
autres caracteres sont ceux des Aphrophores. Du gree /^-'V'Sr, 
coquille d'oeuf." 



Doering: Lepyronia quadrangularis. 517 

The genus Leypronia is separated by Ball from the other genera 
of Aphrophorinse by these characteristics: Anterior margin of ver- 
tex between front and eyes sharp ; ocelli nearly equally distant from 
eyes and each other; rostrum short, not exceeding middle coxae; 
anterior margin of pronotum rounded; corium without terminal 
membrane ; whole upper sm-face densely pubescent, hiding sculptur- 
ing and venation. 

KEY TO SPECIES * 

A. Margins of the vertex regularly rounding to the obtuse tip; elytra 
islightly angularly inflated, nearly twice longer than their combined 
width (folded), graj-ish, testaceous, with a distinct V on each elytron. 

quadrangularis (Say) . 

AA. Margins of vertex straight or concave, the tip slightly produced; elytra 
inflated, no more than one-half longer than their combined width 
(folded). 
B. Small, testaceous, rather narrow; the vertex broad and short, 
shorter or only equaling the pronotum in length; apex of elytra 
broadly subhyaline. angulifera (Uhl). 

BB. Large, nearly uniform grayish, general form globose; vertex longer 
than pronotum. gibbosa (Ball) . 

The original description of Say is given herewith: 

Brownish-cinereous elytra with two oblique brown bands confluent at the 
outer margin; beneath black; feet annulate with pale. Body brownish 
cinereous, covered with dense, minute hairs; head obsoletely spotted; eyes 
fuscous, a pale longitudinal line on the middle, in which is a brown central 
line; stemmata indistinct, black; thorax emarginate at the anterior angles 
for the reception of the eyes, and deeply emarginate behind for the reception 
of the scutel; a double series of obsolete, indented spots before; scutel, tip 
and basal angles acute; hemelytra pale brownish cinereous; an oblique black- 
brown fascia from inner basal angle is confluent at the middle of the exterior 
margin, with an oblique fascia, which terminates near the sutural tip; tip with 
a small blackish curve; region of humerus dusky, beneath black; feet black; 
thighs annulate with pale; posterior pair of tibiae pale, armed with two 
robust spines behind and numerous small ones at the tip; posterior tarsi 
armed with spinules at the tips of the first and second joints beneath; abdomen 
black; tail pale beneath. 

The following is a technical description: 

Form. Length: 9 6.4 to 7.2 mm.; 5 6 to 6.8 mm. Width: 5 3 to 3.2 
mm.; $ 2.4 to 3 mm. 

Vertex flat or depressed; length and width about equal, as long as pronotum, 
margins rounding to a bhmt apex; tylus large, parallel margined, nearly one- 
half length of vertex; eyes level with vertex, distended below; front somewhat 
inflated, rising gradually from the sides, margins convex, slightly longer than 
wide; clypeus only shghtly inflated, twice as long as wide; pronotum flat, twice 
as wide as long, slightl}' emarginate anteriorly with transverse rows of im- 

*BaIl, E. D. A Review of the CereopidsB of North America North of Mexico. Rept. la. 
Acad. Science, 1898. 



518 The University Science Bulletin. 

pressions behind margins; lateral margins nearly parallel, longer than short 
diameter of eye; elytra not quite twice as long as wide, outer margin flaring, 
apex angulate; abdomen broadly triangular, about as wide as long, margins 
and apex greatly exceeded by wings. 

Color. Vertex and tylus mottled testaceous brown; minute yellow line on 
margins; eyes deep fuscous; face uniform testaceous brown; pronotum reddish 
brown at base, fading into brown cinerous; elytra grayish or tawny brown, a 
patch at the base, another at apex, an oblique band from tip of scutellum to 
a point beyond middle of costa, another from point of claws, meeting this on 
costa and forming a V on each elytron, brownish fuscous; abdomen blackish 
brown, apex paler; legs testaceous brown, annulate with pale, spines on pos- 
terior leg very black at tips. 

LIFE HISTORY. 

HISTORY. 

Life history notes on the Cercopidae are rather limited. Osborn 
(1916) made some valuable studies of the life histories of Maine 
froghoppers, but gave no complete history, including the egg stage 
and five nymphal stages, of any one species. He figures and de- 
scribes three instars of Lepyronia quadrangularis. In 1921 Garman 
published the life history of Philcenus lineatus (Linn.), wherein he 
describes the egg and four instars. Later Barber and Ellis (1922) 
described the oviposition of three species, Philcenus lineatus (Linn.) 
and Philaroiiia bilineata (Say.) The most complete study of a cer- 
copid life history is that of a foreign form, Tomaspis varia, which is 
a pest of sugar cane in Trinidad (Urich, 1913). Garman (1923) 
gives a complete description of the life history of Clastoptera obtusa 
(Say) and a brief description of the egg stage of Lepyronia quad- 
rangularis (Say). 

DISTRIBUTION. 

Lepyronia quadrangularis (Say), according to the Snow collec- 
tion, has been taken from four counties in Kansas, namely, Chero- 
kee, Douglas, Neosho, and Doniphan. It is also recorded here from 
Atherton, Mo. Ball (1898) lists it from Ontario, New Hampshire, 
New York, Pennsylvania, Connecticut, District of Columbia, Mary- 
land, West Virginia, Georgia, Florida, Mississippi, Ohio, Iowa, 
South Dakota, Nebraska, Colorado and Texas. Van Duzee lists it 
from Muskoka lake district of Canada, Lake Temagami, Ontario 
and Quinze lake region. Comstock says it is one of the most com- 
mon spittle bugs of eastern United States. 



Doering: Lepyronia quadrangularis. 519 

HABITAT. 

Lepyronia quadrangularis is usually associated with weed patches, 
and therefore might truly be called a weed insect. Yet its habitat is 
not quite so general as this would imply. Neither adults nor 
nymphs can be taken on every weed patch one runs across in col- 
lecting, but seems to be found only in particular places. The se- 
lected spots usually occur near woods or thickets. It has been taken 
in thickets where weeds have sprung up in open spaces, by the road- 
sides adjoining woods or thickets, and on the weeds growing at the 
edge of a wheat field and on the wheat itself. They were found in 
most abundance, however, in a weed patch occupying half of a city 
block. This plot offered a variety of host plants and shelter, such 
as dogwood and elm sprouts, and large patches of sweet-clover, rag- 
weed and grasses; in fact, most of the collecting necessary for this 
work was done at this place, and several thousands of spittle insects 
must have been taken here. 

HIBERNATION AND SPRING APPEARANCE. 

Ball (1920) stated that all Cercopidse except one overwinter in 
the egg stage. Having this idea in mind when these studies were 
started, it was expected that the adults soon after emerging would 
mate, the female lay her eggs, and the overwintering form be easily 
obtained. The matter, however, was not as simple as this. During 
the summer of 1921 adults were observed in the field until August, 
but no mating or oviposition took place. The following spring, 
toward the latter part of April, eggs were sought in the field, but 
none could be found. On April 29 the first instars were found. At 
this time Mr. C. H. Curran gave the writer two adult females which 
he had collected on April 1 and April 4. He stated that toward the 
latter part of March and April adults had been quite abundant. 
This evidence seemed to point to the supposition that the adults 
might overwinter. Throughout the summer adults were closely ob- 
served in both the field and laboratory. During August and Sep- 
tember adults were very abundant, an average of fifty being taken 
in an hour's collecting. On September 27, seventy-five adults were 
taken; on October 9, sixty; and October 19, thirty. In one of the 
outside cages four adults were observed on November 23. These 
late occurrences were due, no doubt, to the very late season, which 
prolonged vegetation as well as insect life. 

In the late fall, after mating had taken place, an experiment was 
attempted for the purpose of obtaining a premature oviposition. 



520 The University Science Bulletin. 

Twenty adults were placed in a glass jar containing sterilized leaves 
and twigs and covered by cheese cloth. The jar was then placed 
outdoors, surrounded by ice except for a small breathing space at 
the top and kept in this condition for eight days. After a few days 
the adults became very sluggish, scarcely moving when disturbed 
except for a slight motion of the legs. When the eight days had 
elapsed they were brought into a warm room and placed in a cage in 
which green plants were growing. Nine out of twenty insects sur- 
vived, but no oviposition occurred. Although not fulfilling its orig- 
inal purpose, the experiment seemed to show that the adults probably 
overwinter. On December 23 Mr. Beamer collected a female of this 
species while sifting leaves in Cherokee county. Four overwintering 
females were taken on April 18 and 19 of this year. The exact place 
of hibernation is difficult to determine. To date no adults have been 
obtained from the outside cage. They probably overwinter deep 
down under the leaves and matted grass in the cracks and crevices 
of the ground.* 

CLIMATOLOGICAL DATA. 

As was stated in the foregoing paragraphs, overwintering females 
were taken on April 1 and April 4 in the spring of 1922, and per- 
haps others could have been obtained earlier, while in 1923 the 
first ones were taken on April 19, making a total difference of 
seventeen days. This great variation can easily be explained by 
the difference in temperature of the two years. A comparison of 
the two springs is given in the following table: 

Mean Departure. Lowest Highest Greatest 

19S2. for month, from normal. temperature. temperature, daily range. 

February 34.2° +1.47 2° on 13th 75° on 21st 39o 

March 44.5° +1.4 10° on 1st 78° on 23d 36° 

April 56.3° +1.7 31° on 1st 82° on 6th 38° 

ms. 

February 30.3° — 0.54 —2° 61° 38° 

March 40.74° —2.03 ^1° 80° 44° 

From a study of the table it is readily seen that the spring of 
1922 was above normal, while this spring is below normal, which 
accounts for the great difference in the dates of spring emergence. 

SEASONAL history AND SUMMARY OF LIFE HISTORY. 

A brief seasonal history and summary of life history is as fol- 
lows: The insect spends the greater part of the year in the adult 
form, covering a period of about ten months, and the other two 
months are spent in the egg and nymphal stages. Mating takes 
place in the fall; the females overwinter, emerging in the spring 
along in April. A few days after emergence the eggs are laid. 



Doering: Lepyronia qladraxuularis. 521 

The egg stage probably lasts two weeks, followed by a six weeks' 
nymphal stage. The adults appear in June and feed all summer 
and fall until after mating, when the males probably die and the 
females go into winter quarters. 

OVIPOSITION. 

Females collected in the spring were confined on small Solidago 
and sweet-clover plants under lamp chimneys. Three females col- 
lected on April 18 died on April 23. Two of the females apparently 
laid no eggs. Thirty-eight eggs were found in the abdomen of one 
and thirty-three in the other. The third female laid in all nine 
eggs, one of them being found alone in one leaf and the other eight 
in a group in another leaf. The latter were laid in a row along the 
slender petiole of the Solidago leaf. Both leaves in which the eggs 
were inserted were brown and withered and were found at the base 
of the plant. This same female was dissected and six eggs were 
found in the abdomen. Another female collected on April 19 died 
April 23 without laying any eggs. Forty-five eggs were found in 
her abdomen. Two other females were taken on April 24, from one 
of which, at this time of writing, only one egg has been obtained. 
It too was inserted in a partially dead leaf. 

The eggs are inserted in the plant tissue. A longitudinal slit is 
made in the leaf and the egg deeply inserted, so that it makes a 
slight bulge in the leaf on the opposite side. The slit appears to 
be plugged with a whitish substance, which probably is part of the 
plant tissue. 

NUMBER OF INSTARS. 

The nymph passes through five nymphal stages, each stage differ- 
ing from the others somewhat, both in structural detail and color. 

LENGTH OF THE STAGES. 

The length of the various stages was difficult to determine, due to 
the difficulty in rearing the insects. The work accomplished dur- 
ing the summer of 1921 was of little value. It was started with 
nymphs of the third, fourth and fifth instars. These were brought 
into the laboratory on large cuttings of host plants, which w^ere 
placed in water and covered by lamp chimneys. The intention was 
to change the spittle insects to fresh plants as quickly as the original 
ones wilted. Apparently there was not enough plant juice to supply 
the amount of fluid for so many new masses of spittle necessitated 
by the changes. A few nymphs, however, were carried through to 
adults. In the spring of 1922, having obtained the very small first 



522 



The University Science Bulletin. 



instars, and using a different method of rearing, better results were 
obtained. Various host plants were planted in small flower pots, 
and the insects were confined on these under lamp chimneys. This 
was an improvement over the first method, although not entirely- 
satisfactory. By having the living plant a more steady flow of sap 
could be obtained, so that the insect, once established was able to 
proceed without interruptions. The chief difficulties to cope with 
were in getting the very restless first instars settled, and the fact 
that the plants became stunted and sickly under glass 'chimneys. 
Wire cages would have been more satisfactory for the rearing of the 
later nymphal stages, but would have been too open for the first 
instar. From some fifty-odd groups of experiments, only seven 
first-instar nymphs were carried through to adults. Other reliable 
data for the various stages were obtained piecemeal by counting the 
length of time from molt to molt. The rearings were carried on on 
a back porch, so that the temperature was practically the same as 
in the field. 

First-instar nymphs taken in the field remained the following 
number of days in this stage : 



April 29 to May 5 7 days 

May 2 to May 9 7 days 

May 3 to May 10 7 days 

May 4 to May 14 10 days 

May 6 to May 17 U days 

May 12 to May 17 5 days 



May 12 to May 21 9 days 

May 12 to May 19 7 days 

May 12 to May 20 8 days 

May 12 to May 21 9 days 

May 12 to May 14 2 days 

May 1 1 to May 21 10 days 



This gives a range of from two to thirteen days for the instar. 
However, these data are not sufficiently accurate, since the nymphs 
were not obtained directly from the egg, but were picked up pro- 
miscuously in the field. Some of them appeared to have just 
hatched. The average time is probably ten or eleven days. 
Records for the second instar are as follows: 

May 21 to May 27 6 days 

May 25 to June 3 9 days 

May 22 to May 24 2 days 

May 22 to May 27 5 days 

May 22 to May 28 6 days 

May 24 to May 31 7 days 

June 26 to June 30 4 days 



May 17 to May 25 8 days 

May 17 to May 22 5 days 

May 21 to May 27 6 days 

May 19 to May 25 6 days 

May 19 to May 26 7 days 

May 21 to May 29 8 days 

May 14 to May 21 7 days 

These are accurate counts taken from molt to molt. The range 
is from two to nine days and the average length seems to be six or 
seven days. 



Doering: Lepyronia quadrangularis. 



523 



Tliird-instar nymphs lived from — 

June 3 to Juno 9 6 days 

May 24 to June 5 12 days 

May 28 to June 8 11 days 



May 25 to May 31 6 days 

May 22 to May 29 7 days 

May 27 to June 6 10 days 

May 25 to June 1 7 days 

May 26 to June 3 8 days 

May 29 to June 6 8 days 

Mav 21 to May 30 9 davs 



May 31 to June 13 13 days 

May 27 to June 4 8 days 

June 30 to July 6 6 days 



These are also counted from molt to molt, 
six to thirteen. The average is eight. 
The fourth stage lasted from — 

May 31 to June 8 8 days 

June 6 to July 1 25 days 

June 1 to June 11 10 days 

June 3 to June 12 9 days 

June 6 to June 16 10 days 

May 30 to June 9 10 days 

June 9 to June 18 9 davs 



The range is from 



June 5 to June 14 9 days 

•June 13 to July 7 24 days 

May 31 to June 12 12 days 

June 5 to June 13 8 days 

June 6 to June 13 7 daj's 

June 8 to June 14 6 days 

June 1 to June 25 24 days 



The range is from six to twenty-five and the average is between 

nine and ten. 

The fifth stage lasted from — 

June 8 to June 19 11 days 

June 1 to July 15 14 days 

June 11 to June 26 15 days 

June 12 to June 28 16 days 

June 16 to July 8 22 days 

June 9 to June 23 14 days 



June 18 to June 29 11 days 

June 14 to June 24 10 days 

July 7 to July 20 13 days 

June 12 to June 30 18 days 

June 14 to June 25 11 days 

June 25 to July 9 14 daj'-s 

The range is from ten to twenty-two and the average between 
eleven and fifteen. 

The seven complete histories from first instar to adult are as 
follows : 

Number. 

1 

2 

3 

4 

5 

6. 

7 



First. 


Second. 


Third. 


Fourth. 


Fifth. 


Totals. 


11 


8 


6 


8 


11 


44 


9 


6 


10 


25 


14 


89 


7 


6 


7 


10 


15 


45 


7 


7 


8 


9 


16 


47 


9 


8 


8 


10 


22 


57 


2 


7 


6 


10 


14 


39 


13 


9 


6 


9 


11 


48 



From the foregoing data it is apparent that there is a wide varia- 
tion in the length of any stage, which is probably due to the nature 
and condition of the food plant and the abundance of the sap. 

Field observations also show a wide variation in the length of the 
stages. In 1922 the first-instar nymphs were taken on April 29 and 



524 The University Science Bulletin. 

the last one on June 23, while they reached their maximum abun- 
dance between May 11 and May 16. Second-instar nymphs ap- 
peared May 4, reached their maximum number about May 18 to 
May 27, and disappeared around June 20. Third instars were first 
taken about May 11, were most abundant from May 29 to June 3, 
while fourth instars appeared about May 16 and reached the maxi- 
mum from June 7 to June 10. The first fifth instars were taken on 
June 2. They were most abundant from June 13 to June 20 and 
began to thin out by June 27, the last two being taken on July 4. 
The average total of days for the nymphal stages is perhaps forty- 
five days. 

HABITS OF NYMPHS. 

• 

Perhaps the first thing of interest to note about the nymphs is 
their restlessness in the laboratory and their agility in moving 
around. Since they are always found in spittle masses, it is only 
natural to think of them as being rather inactive, helpless larvae, 
but such does not appear to be the case. The first-instar nymphs 
especially are most active. In fact one of the greatest difficulties 
to overcome in the rearing of these insects was the continual moving 
of the first instars. When collecting, if the larva were brought away 
undisturbed in their spittle masses they usually were found missing 
on arriving at the laboratory. A satisfactory method of taking 
them is to wrap the plants in an improvised envelope of newspaper 
and carry them away in this fashion. When the nymphs are trans- 
ferred to new plants it takes them some time to become settled. 
They ramble aimlessly over the plant, seemingly looking for the 
most favorable feeding ground. In one instance a nymph was 
observed which wandered the entire length of the stem and back 
again, then out on the leaf, where it rambled all over the surface, 
and even hung poised in the air, walking on the extreme margin of 
the leaf. During this wandering period they repeatedly fall off the 
plant on the dirt, where, if they happen to light on their backs, they 
struggle to regain their feet, and usually perish in the attempt. They 
dry up very quickly if not living in their spittle masses. Many times 
they crawl from the plant to the ground and reach the surface of the 
glass globe, where ensues a struggle to climb up the slick glass. 
Several times during their wanderings the first instars have dis- 
appeared entirely, crawling through two thicknesses of cheesecloth. 
This restlessness of the nymphs is apt to be incited in two ways, 
namely, by disturbing them in the spittle mass and because of lack 
of juice from the plant. While looking for molted skins it was 



Doering: Lepyronia quadrangularis. 525 

usually necessary to shove the insect around in the spittle mass in 
order to see the skin, and this often disturbed the nymph so much 
that it would immediately hunt a new place. 

Lepyronia nymphs have a comical appearance while walking over 
the plant. Their legs are long and they walk with their bodies lifted 
high in the air. Sometimes they walk exceedingly fast, but at other 
times they merely creep along. Occasionally the nymph extends 
its abdomen in the air at right angles to its body, first expanding it 
and then contracting it in a telescopic manner; it does this even 
while it walks. 

The gregariousness of spittle insects is pkinly evident, although 
it probably is due more to chance than instinct. A probable ex- 
planation is that in their roaming over the plants they encounter 
other spittle masses, and it is much easier to stay in this than to 
make a new mass. In many cases three or four instars have been 
found together in the same spittle mass. For this reason, toward 
the last of the season the first instars are not so readily seen unless 
each spittle mass is examined, since they are found deeply imbedded 
in the spittle mass made by larger nymphs. On one stalk of Ambro- 
sia trifida (horseweed) three inches of solid spittle were found ex- 
tending all around the stem. In the mass there were thirty-one in- 
sects, of which sixteen were fourth instars, ten were third instars, 
and five 'were fifth instars. Fifteen molted skins were found in it 
also. On another plant were found three large masses of fourth and 
fifth instars, which were packed so closely together that the spittle 
scarcely covered them. Spittle masses were very large and abun- 
dant on elm sprouts; one stalk bore a mass of spittle which extended 
four inches along the stem ; another branch bore eleven masses. On 
another host plant two large masses of spittle were found, one mass 
containing six nymphs and the other six or ten. On June 13, on a 
single plant of Ambrosia trifida a spittle mass twelve inches long 
was found, which contained sixty-eight or more nymphs. The spit- 
tle was white and foamy, but was barely enough to cover the 
nymphs, since the form of their bodies could be plainly seen 
through it. 

In the literature ii is often stated that the nymph lives and molts 
in the same mass of spittle until the adult form is reached. Com- 
stock (1895), in his brief discussion of Cercopidae, states that it had 
been asserted that they undergo all their changes in the spittle mass, 
Girault (1904) says of Aphrophora parallela (Say), that they seldom 
move unless disturbed, and Garman (1921), in his work with Phil- 



526 The University Science Bulletin. 

(Buus lineatus (Linn.), states that during nymphal life the bug may 
construct several balls, but that there usually is little migration 
after the first mass is formed. Kershaw (1914) , on the other hand, 
is of the opinion that the nymphs of Tomaspis saccharina usually, 
but not always, leave the spittle mass after a molt. 

In the study of Lepyronia quadrangularis an experiment was at- 
tempted to determine just to what extent these migrations were car- 
ried on in the field. Four goldenrod plants bearing spittle masses 
were marked with white rags about May 12. It was difficult to de- 
termine in what stage the nymphs were at this time, since it was not 
thought best to disturb them, but they appeared to be in the latter 
part of the first stage. On May 16 three plants had second instars 
on them and the fourth had a second- and a fourth-stage nymph in 
one mass. On May 27 these plants were observed again. One plant 
was entirely deserted, but the other three appeared to be all right, 
with the spittle masses in practically the same position. June 2 
found them practically the same, but other weeds were beginning 
to crowd around them so that little data thenceforth could be ob- 
tained. On June 7, however, one or two new masses of spittle were 
found on each of the marked plants, including the deserted one, 
which now bore three masses. This seems to point to the conclusion 
that the nymphs do move in the field even when not disturbed. Fre- 
quently, while collecting, molted skins can be found in deserted 

spittle masses. 

In captivity, at least, as was stated above, the nymphs move con- 
siderably. Following the records of a few reared from first instars 
to adults may illustrate this fact. One first instar collected on May 
6 was placed on a plant and soon formed a spittle mass. On May 
17 it molted to a second instar, remaining in the same mass of spit- 
tle until May 18. On this date it moved farther down on the plant 
and made a new mass. It remained in this mass of spittle until 
June 7, in the meantime having molted on May 23 and May 25. 
After this last migration it formed a new mass of spittle higher up 
on the plant, where it remained until emerging as an adult. Another 
nymph, collected on May 12 and reared to adult, moved only twice, 
both times being between molts and not just after. Still another 
moved three times during its five changes. A fourth specimen, col- 
lected in the second stage, moved six times before emerging as an 
adult, and only once moved directly after molting. From the fore- 
going there appears to be no fixed habit of moving after each molt. 



Doering: Lepyronia quadrangularis. 527 

They probably move from lack of food or because of disturbances, 
and in the field, if they find a favorable place, they may stay in one 
place throughout their period of growth. 

The size of the spittle mass varies for each instar, being in pro- 
portion to the size of the nymphs. The first-instar nymphs, at least 
the recently hatched ones, are covered by a clear drop of fluid with 
very little foam, and for this reason are easily discovered in the 
field. In fact, they were originally discovered by means of these 
clear drops of fluid. When walking through the weeds the writer's 
attention was attracted by these shiny drops of fluid on the vegeta- 
tion, and upon closer examination it was found that they came 
from the spittle nymphs, which were usually found on the under 
sides of the leaves, and for this reason are not easily seen. The 
spittle masses of the other instars are usually foamy and puffy in 
the field. 

When the insect is ready to transform into an adult an interest- 
ing change in the spittle mass takes place. The superficial part of 
the foam dries and stiffens somewhat, so that it forms a roof to a 
closed chamber. Within this chamber the molt occurs, and the adult 
can be seen distinctly in this mass. It usually takes a day for the 
adult to harden sufficiently to emerge. If they are removed from 
the mass too soon they do not gain their full color, but have a yel- 
lowish-tan appearance. When the adult is ready to emerge the 
slightest disturbance causes it to give a strong leap, thus freeing it- 
self from the spittle ball and leaving a round opening in the latter. 
Empty chambers are found quite frequently in the field. 

In captivity all the instars made comparatively little spittle. In 
a good many cases they lived chiefly in the fluid without producing 
many bubbles. When the spittle was produced it usually was just 
sufficient to cover them. The fourth- and fifth-instar nymphs were 
able to make the most spittle, but even these did not reach the 
proportion of those in the field. 

The process of froth-making in the Cercopidae has been a topic of 
discussion for some time. The earliest conception of the spittle 
masses was that they were voided by tree frogs. Fabre (1900), in 
the discussion of froth-making by Aphrophora spumarius, which in 
his picturesque way he calls the "cicadelle," says that the peasants 
of France give another name to this substance. They call it "cuckoo 
spit," because the little balls occur at the time of year when this 
bird returns from its migrations. The early entomologists assigned 
the spittle to its correct cause, but they thought that the foam 



528 The University Science Bulletin. 

was exuded from the arms of the little nymphs. This viewpoint was 
summarized by Harris (1862) as follows: "Here may be arranged 
the singular insects called froghoppers, Cercopidse, which pass their 
whole lives on plants, on the stems of which their eggs are laid in 
the autumn. The following summer they are hatched and the young 
immediately perforate the bark with their beaks and begin to im- 
bibe the sap. They take in such quantities of this that it oozes out 
of their bodies continually in the form of little bubbles, which soon 
completely cover up the insects." 

In 1900 several papers on froth formation were published by 
European and American writers. Morse (1900) is generally given 
the credit for discovering that the insect emits a liquid only and 
later enfolds air bubbles in the liquid. His original account really 
appeared in 1875 in his "Elementary Zoology." Fabre (1900), 
Gruner (1900), and Sulc (1900) concluded the same thing, but they 
all differ as to the method by which the result is obtained. 

According to Morse (1900) a clear fluid is emitted by the nymph, 
which flows over the entire body and fills up the crevices between 
the legs. Next the insect extends the abdomen out of the fluid, 
opens the posterior segments like claspers, grasps a bubble of air, 
and then turns the abdomen under the fluid, allowing the inclosed 
air to escape. According to him, the movements go on at the rate 
of seventy to eighty times a minute and thirty to forty bubbles 
were made in a half hour. He says that the claspers seem to be the 
tergal portions of the ninth segment. 

Fabre (1900) describes a similar apparatus. According to him, 
the insect has a special device, which is composed- of the two pleural 
lobes of the ninth segment, acting as claspers for grasping air, and 
a pocket, formed by these lobes, which serves as a container for 
air. From a caudal view of the abdomen, when the two pleural 
lobes are drawn apart, a y-shaped opening in the pocket is produced, 
or, in other words, the expanding and contracting of these lobes 
opens and closes the pocket. In producing the bubbles the tip of the 
abdomen is thrust out of the liquid, the pleural lobes spread apart, 
letting air into the pocket, then close together again, and at the 
same time the abdomen is pulled under the fluid. At this point the 
pocket, being flexible, contracts, and thus forces air out of the 
pocket, forming a bubble in the viscid fluid. 

The explanation of bubble formation advanced by Gruner (1900) 
is similar to that of Fabre's in that he too describes a pocketlike 
cavity and two terminal clasping plates. He maintains, however, 



Doering: Lepyronia quadrangllaris. 529 

that the air for the bubbles is supplied from the tracheae. The fol- 
lowing is his account of the process: ''Soon after the larvae have 
fastened themselves head downward on the plant and have imbibed 
some of the sap. the terminal portion of the abdomen rythmically 
contracts so that the fluid from the anus is exuded and flows into the 
cavity or pocket. The insect being head downward, the fluid flows 
into the pocket, where it becomes mixed with air coming from the 
last few pairs of spiracles. Bubbles are thus produced in the pocket 
by the contraction and expansion of the tergal plates." 

Sulc (1910) describes a still different apparatus, namely the air 
canal. He says that the "pochette" of Fabre and the "tasche" of 
Gruner is not present, but that the special device is the air canal 
(pi. LXII, figs. 6 and 7). This air canal or channel is formed by the 
tergal pads or plates. Beginning with the fourth segment and ex- 
tending to the ninth, the plates are prominent and are capable of 
touching on their median margins. The plates of the first and 
second segments are short and widely separated from each other 
The tergal plates of the third abdominal segment are triangular, so 
that only their posterior medial margins can touch each other. 
Arising from the median portion of the third sternite between the 
two plates there appears a special triangular, caudad-projecting 
protuberance (pi. LXII, fig. 6), which serves to close the channel. 
Thus the tergal plates form a channel which extends directly from 
the ninth segment to the rniddle of the third. At this point it be- 
comes Y-shaped, dividing into two smaller channels, which continue 
to the right and left until the hind margins of the thorax are reached. 
When the cercopid is submerged in the spittle the tergal pads are 
pressed firmly together by means of the contraction of strong 
muscles, and the tip of the abdomen just reaches the surface of the 
spittle. According to Sulc, the froghopper nymph can be compared 
to water-dwelling insects with open, tracheal systems, only devi- 
ating from all hitherto know^n examples by having this special de- 
vice of the air channel. When fresh air is desired the tip of the 
abdomen is thrust out of the spittle, the air channel opens and the 
air enters. Immediately the air channel is compressed and at this 
moment a bubble is released in the spittle mass. 

The writer's own observations were made before reading the 
above descriptions. It was very difficult to view this process at all, 
because of the difficulty of getting the nymphs to settle on the cut 
plants, and when they made the foam on living plants it was almost 
impossible to focus the microscope on them. Most of the observa- 



530 The University Science Bulletin. 

tions were interrupted before much data could be obtained. The 
longest observation was that on a fourth instar, which started mak- 
ing spittle on a large leaf which could easily be placed under the 
microscope. It started making spittle at 9:40 a. m. At this time 
the abdomen was extended in the air, and from the slitlike opening 
in the anus could be seen to exude a clear fluid, which flowed down 
beneath the body in the ventral channel or air canal of Sulc. A 
second or so was spent in ejecting this fluid, and then the insect 
began dipping the abdomen in and out of this fluid, each time pro- 
ducing a bubble. It did this for ten times, and then the abdomen 
was lifted high in the air twice and the terminal plates were spread 
far apart. When a number of bubbles was produced the abdomen 
was pulled beneath them and then brought up through them so that 
they were separated and pushed on each side of it. When reaching 
after air the plates were spread far apart, but at the surface of the 
spittle they came together. The farther out the abdomen was ex- 
tended and the slower the movements the larger the bubbles were. 
The abdomen was next pulled down deeply in the spittle and a few 
small bubbles were produced. Again the nymph rested for a minute 
or so, while more fluid flowed from the anus. At times the nymph 
kicked rapi'dly with its two front pair of legs, making the bubbles 
go to the side, and during the whole performance one front leg was 
constantly kicking so that gradually the bubbles were worked head- 
ward. Again it made bubbles, rested for three and one-half minutes, 
while the body was straightened out until the spittle was all around 
the side and partly on the side of the head. Another cessation of 
bubble-making lasted for six minutes. Later the nymph lay almost 
on one side with the bubbles entirely on the other side. Next more 
fluid was ejected until the. body was entirely surrounded and the 
bubbles pushed entirely to the edge of the fluid mass. A rest of ten 
minutes followed. At 11:10 it had not ceased making spittle, and 
the insect was still visible, although the entire body was covered by 
a thin sheet of foam. At 11:45 there was considerable more spittle 
and the nymph was still working. 

Another nymph was watched. It started making spittle at 8:20, 
and in fifteen minutes had enough spittle to make a thin covering 
for its body. 

From a study of the nymphs of Lepyronia it appears that Sulc 
has described the process more accurately than any of the others. 
His description seems to differ only in extent from that of Fabre and 
Morse, both of which consider the two tergal plates of the ninth 



Doering: Lei'YRonia quadkangularis. 531 

segment of the abdomen as being the responsible machinery. Since 
these are the terminal plates, they naturally are the most prom- 
inent and mider low magnification appear to do most of the work. 
From a translation of Sulc's work, it seems that he does not attrib- 
ute the bubbles to the work of merely the last pair of plates, but 
to all nine pairs, which come together and form the air channel. He 
thus makes respiration and bubble-formation a simultaneous action. 
From observations of Lepyronia he is apparently right, since the 
writer has seen six pairs of plates in action very distinctly. 

COMPOSITION OF THE FROTH. 

The composition of the froth of spittle insects has been studied 
by different writers. They all have noted the viscid quality of the 
fluid, which they assert is necessary to maintain the frothy condition. 
Morse (1900) found that v/hen nymphs were placed in water they 
immediately began clutching the air and making bubbles. The 
bubbles disappeared, however, as soon as made, for the clear water 
did not preserve them. Besides this viscidity, the spittle has a cer- 
tain insolubility in water and alcohol. When collecting immediately 
after showers and heavy rains the spittle masses are found intact, 
even those which were in exposed places. In alcohol the spittle 
seems to coagulate in a stringy mass, which clings to the feet and 
body and is hard to pull away. Garman (1921) says that it is 
readily soluble in sodium hydroxide, has no reaction to iodine, al- 
though probably containing some sugar, and that the albuminous 
substance is not coagulated with heat. Gruner (1900) after several 
experiments comes to the conclusion that 94.565 per cent of the 
spittle fluid is water, 3.827 per cent organic substance, and 1.607 per 
cent inorganic salts. The spittle mass of Lepyronia can easily be 
told in the field from that of a Clastoptera by its composition. The 
spittle of the former contains more and larger bubbles and resembles 
the beaten white of egg, while the spittle of Clastoptera contains 
much smaller bubbles, which seem to be all on the outside of the 
mass, while the center of the mass appears to be a clear, gelatinous 
material. Kershaw (1914) states that the spittle of the nymph 
appears to be a mucin or mucinoid. He found that it granulates 
with subacetate of lead and stains deeply with methylene blue in 
glycerin and alcohol. Furthermore, the fluid appears to have every 
substance excreted from the anus, such as calcium oxalate, uric acid, 
leucine pellates and urates, potassium and sodium chlorides. 



532 The University Science Bulletin. 

LONGEVITY OF ADULTS. 

In 1921 the first adults were collected June 14 and the last ones 
on July 23. The latter date does not represent their last appearance, 
but merely the time when collecting trips ceased. In the spring of 
1923 Mr. C. H. Curran collected specimens on April 1 and April 4, 
which was practically the first indication that they might overwinter 
as adults. By the first part of May adults were no more in evidence. 
Adults hatching from eggs in the spring began to appear on June 13, 
becoming quite numerous by June 19. The rest of the summer and 
fall adults could be taken in great numbers, sometimes averaging 
seventy-five in one hour's collecting. The last date in the fall was 
November 23. In the spring of 1923 adult females were first taken 
on April 19. From these facts it appears that the insects, at least 
the female, spends the greater part of the year in the adult form — 
from the middle of June until about the middle of April, or a period 
of ten months. 

HABITS OF ADULTS. 

The adult bug is a very sluggish insect. It sits for hours on the 
stems and leaves of plants, feeding continually. Many were found 
on thistle plants, where they could be easily watched. Some were 
under observation for two hours, during which time they never 
changed position. While feeding they emit a clear fluid or honey 
dew, which falls from beneath them in large drops. One individual 
was watched for an hour, during which time 200 drops of honeydew 
were emitted. While feeding they usually sit with their legs folded 
snugly beneath the wings. When walking they seem to spread the 
two front pairs of legs out to the side, propelling themselves along 
by them, but the hind pair are held straight beneath the wings and 
dragged along behind, evidently only of use in jumping. When 
disturbed froghoppers give a powerful leap, and for this reason are 
hard to collect. The best method of collecting them is by sweeping 
and then holding a large-sized test-tube in front of them in such a 
position that they will hop into the tube, since they always jump 
in a forward direction. 

MATING. 

Mating takes place in the late fall. On September 27 two pairs 
were found mating in a cage in the insectary. On September 28 
one pair was caught in the field while mating and two pairs in an 
inside cage. A day later two more pairs mated, and on September 30 
another pair and on October 2 still another, making nine pairs in 
all. After this the males probably die and the females go into 
winter quarters. 



DoERixc;: Lhpyronia quadrangularis. 533 

FOOD HABITS. 

The nymphs and adults of this cercopid have a wide range of 
food phmts. The nymphs have been found feeding on sixty-two 
species of phmts, which are as foHows: 

Actinonioris altcrnifolia Compositse. 

Agrimonia {iyi'Posei)aIa (agrimony) Rosacea). 

Ambrosia trifida (horseweed) Compositse. 

Ambrosia artemisiifolia (ragweed) Compositse. 

Andropogan furratus Gramineae. 

Apocynuiu caniiabinuni (dogbane) Apocynaceae. 

Apocynvun luibescens Apocynaceae. 

Asclepias syriaca (common milkweed) Asclepiadacese. 

Asclepia.s tuberosa (butterfly weed) Asclepiadaceae. 

Asclepias verticillata Asclepiadaceae. 

Aster paniculatum Compositae. 

Aster salicifolius Compositae. 

Bromii^ secalinus (chess or cheat) Gramineae. 

Chenopodium album (lamb's-quarters) Chenopodiaceae. 

Cirsium altissimum (thistle) Compositae. 

Cornus baileyi (dogwood) Comaceae. 

Erigeron canadensis (butterweed) Compositae. 

Erigeron ramosus (daisj^ fleabane) Compositae. 

Eupatorium altissimum Compositae. 

Galium aparine var. vaillantii (cleavers) Rubiacese. 

Geum canadense (avens) Rosaceae. 

Geum strictum (avens) Rosaceae. 

Helianthus grosserratus (?) (sunflower) Compositae. 

Helianthus petiolaris Compositae. 

Hehopsis scabra (oxeye) Compositae. 

Juglans nigra (black walnut) Juglandaceae. 

Lactuca canadensis (wild lettuce) Compositae. 

Lactuca pulchella (?) Compositae. 

Lactuca scariola Compositae. 

Lactuca spicata Compositae. 

Lycopus americanus (?) (water horehound) Labiatae. 

Melilotus alba (sweet-clover) Leguminosae. 

Morus rubra (red mulberry) Urticaceae. 

Osmorhiza claytoni (sweet cicely) Umbelliferae. 

Panicum virgatum Gramineae. 

Panicum wilcoxianum Gramineae. 

Phleum pratense (timothy) Gramineae. 

Plantago rugelii (plantain) Plantaginaceae. 

Prunella vulgaris (heal-all or carpenter weed) . . . Labiatae. 

Rhus glabra (smooth sumac) Anacardiacese. 

Robinia pseudo-acacia (black locust) Leguminosae. 

Rosa setigera (wild rose) Rosaceae. 

Rubus canadensis Rosaceae. 

Rubus occidentalis (black raspberrj) Rosaceae. 



534 The University Science Bulletin. 

Rudbeckia fulgida (coneflower) CompositEe. 

Ruellia ciliosa Acanthacese. 

Ruellia parvifiora Acanthacese. 

Sanicula canadense Umbelliferge. 

Sanicula marilandica (black snakeroot) Umbelliferae. 

Solidago altissima (goldenrod) Compoeitae. 

Solidago rigida Compositse. 

Sphenopholis obtusata Gramineae. 

Stachys palustris (hedge uettle) Labiatse. 

Symphoricarpos orbiculatus (coralbeiTy) Caprifohaceae. 

Taraxacum officinale (dandelion) Compositse. 

Triadenum virginicum (marsh St. John's wort) . . Hypericaceae. 

Tridens flavus (tall redtop grass) . Gramineae. 

Triticum vulgare (wheat) Gramineae. 

Ulmus fulva (slippery elm) Urticacese. 

Urtica gracilis Urticaceae. 

Verbena urticaefolia Verbenaceae. 

Vernonia baldwini (ironweed) Compositae. 

Among this list of plants there are nineteen families of plants 
represented. Tiie families containing the most species are the Com- 
positae, Graminese and Rosacese, which have twenty, nine and five 
species, respectively. In early spring the first nymphs were found 
in one small plot of Solidago plants. Later they were found on 
other groups of plants, such as aster, grasses and giant horseweed 
{Amhrosia trifida) . Toward the last part of the nymphal season 
the most popular single food plant, perhaps, was Amhrosia artemsii- 
folia, or ragweed. It was chiefly on Ambrosia trifida and Cirsium 
altissimum, however, that the large masses of spittle containing so 
many nymphs were found. 

The spittle masses are usually found along the plant stems, 
although with broad-leafed varieties of plants, such as plantain and 
elm, they are also found on the back of the leaves. 

Not only did the nymphs collectively show a remarkable diversity 
of choice of food plants, but each individual nymph seems to have 
no restriction to any particular plant. Nymphs feeding on thistle 
in the field could be brought into the laboratory and reared on rag- 
weed. In the cages where several species of plants were growing 
together, such as ragweed and lamb's-quarters, nymphs have been 
observed to migrate from one to another several times during their 
development. 

Fabre (1900) discusses this strange disregard, as he calls it, of 
genera and species of plants. He says that it would be hard to 
make a list of the plants in his neighborhood which have been lack- 
ing in spittle. With a brush he picked up nymphs feeding on one 



Doering: Lepyronia qtjadrangularis. 535 

species of plant and deposited them on a new plant of entirely dif- 
ferent flavor, only to find that the new was accepted without hesita- 
tion. He found that the insect could easily be transferred from the 
bean, a plant of mild flavor, to the spicy euphorbia, and back again. 
It also could be transferred from such pepper plants as Arum itali- 
cum, of which it takes only a small portion of the leaf to burn the 
lips, to the perfumed niarum and common dandelion. In order to 
find the explanation of this Fabre carried on the following experi- 
ments. He first discovered that when he punctured a euphorbia 
plant with a small instrument, the milky, poisonous sap oozed forth, 
but that when the beak of the insect was pushed in only a colorless, 
neutral fluid was obtained. In fact, the nymph soon perished in the 
milk of the euphorbia, because of its caustic properties. He there- 
fore concludes that the siphon of the cicada, by a selection which 
should be envied, selects at the bottom of the puncture the substance 
it needs for food, which is the same in all plants, and therefore pro- 
duces the same colorless fluid, no matter what the species of plant is 
on which the nymph is feeding. 

Other observers have found this species feeding on additional host 
plants. Gillette and Baker (1895) took it on Clematis ligusticifolia 
and Carex (Gillette). Lintner (1895) found it very common in 
groves of sugar maple, ''where numbers of them were often met with, 
drowned in vessels of sap." Osborn (1916) took it on Imj)atiens 
biflora. 

NATURAL ENEMIES. 

It is generally conceded that the froth or spittle is a protective 
measure. This is undoubtedly the case, for cercopid nymphs are 
remarkabh* free from parasites or predators. A Syrphus knabi was 
found sucking at a spittle mass for a short while, but soon flew away 
without causing any disturbance. At one time a nabid nymph was 
found sucking the nymph of Clastoptera proteus. The latter was 
found about two inches away from the nearest spittle mass, and 
probably had just left the mass when it was captured by the nabid 
nymph. Three adults of this same species of cercopid were found 
in the web of a small spider. They were all dead. Gruner (1900) 
experimented with red ants and the nymphs of Aphrophora and 
Philcenus. If he placed the nymphs, without any spittle, near the 
ants, the latter immediately pounced on them and began dragging 
them away. However, when he placed grass tufts, bearing nymphs 
in spittle masses, near them, the ants immediately covered the tufts, 



536 The University Science Bulletin. 

but as soon as the mass was reached would stop and turn back. 
Garman (1921) states that the spittle ball offers an ideal medium 
for molds and bacteria, which may sometimes be found in large 
numbers. None of these organisms were found in the spittle balls of 
Lepyronia. Urich (1913) in his studies of Trinidad froghoppers has 
found that they are preyed upon by two birds, a reptile, batrachian, 
spiders, predatory insects, and is the host of two parasitic insects 
and a parasitic worm, which all together play an important part in 
the control of these froghoppers. Williams (1923) reports the larvae 
of Drosophila paradoxa living in the spittle mass of a Clastoptera. 
He states that they undoubtedly kill some of the spittle nymphs. 
Considering the family as a whole, however, it is evident that frog- 
hoppers have few natural enemies, and this probably is due to the 
protection of the viscid spittle material. 

DESCRIPTION OF DIFFERENT STAGES. 

THE EGG. 

Length, 1.2 mm.; width, 0.3 mm. Elongate, slightly curved, tapering to a 
rather sharp point at one end and to a more blunt one at the other end. 
They are white in color, somewhat transparent, and the surface is smooth. 

FIRST INSTAR. 

Size. Average length, 1.53 mm.; width across eyes, 0.456 mm.; width across 
abdomen, 0.549 mm. 

Color. Head and thorax yellowish white washed with brown, the meso- 
thorax and metathorax darker gray-brown. The abdomen is a bright yellowy 
with a pair of oval, orange spots occupying the lateral and part of the dorsal 
and ventral surfaces of the fourth, fifth and sixth segments. The eyes are 
reddish brown and the legs yellowish white washed in brown, especially at 
their bases. 

Structural characteristics. The anterior portion of the head is bulblike, 
distinctly rounded and with the clypeus greatly inflated. The division between 
the front and vertex is indicated by a short line cephalad of each antenna, 
which runs mesad. No ocelli are present. The antennte have nine segments, 
the basal segment a short, stout one, the second and third elongate-stout, and 
the fourth fan-shaped, with the distal five segments ringlike. There is no 
evidence of wing pads from a dorsal view, and only a faint indication from 
the lateral view in the form of a slight caudal extension. The legs have pro- 
portionately greatly elongated coxae and have only two segments in the tarsus, 
a short basal one and a longer distal one. Pleural lobes are not conspicuous. 

SECOND INSTAR. 

Size. Average length, 2.02 mm.; width across eyes, 0-525 mm.; width across 
abdomen, 0.699 mm. 

Color. Body is pale yellowish-white. Head is washed in reddish-brown. 
Prothorax is pale yellowush-white only slightly washed in grayish-brown. 
Mesothorax and metathorax are darker gray-brown. Orange spots on abdomen 
are only faintly visible. 



DoKKiNt;: Lepyronia qtjadrangularis. 537 

Structural characteristics. Practicallj^ the same as in the preceding instar 
The wing: pads show no marked dovolopmont. The mesothorax and meta- 
thorax together are equal in leiigtli to the prothorax. Clypeus is shghtly more 
inflated than in the first instar. 

THIRD INSTAR. 

Size. Average length, 3.75 mm.; width across eyes, 1.05 mm.; width across 
abdomen, 1.33 mm. 

Color. Bodj' is pale whitish-yellow, with considerable more yellow than in 
the second instar. Head and thorax are unchanged in color. 

Structural characteristics. The' wing pads are plainly visible as caudal ex- 
tensions of the lateral angle of the thoracic tergites, the first pair more promi- 
nent than the second. The mesothorax is equal in length to the prothorax 
and about twice the length of the metathorax. Two ocelli are present and are 
black in color. Antennae with nine segments, but the terminal five segments 
are nmch elongated. 

FOURTH INSTAR. 

Size. Average length, 4.48 mm.; width across eyes, 2.57 mm.; width across 
abdomen, 2.17 mm. 

Color. Body pale greenish yellow. Head washed with reddish brown. 
Thorax ^■aries from pale yellow to dark brown on mesothorax and metathorax. 

Structural characteristics. Antennae are practically the same in shape, but 
more elongate. The fii"st pair of wing pads are produced caudad until their 
apices almost reach the apex of the second pair. The exposed portion of the 
second pair is but little longer than that of the first pair. 

FIFTH INSTAR. 

Size. Average length, 6.94 mm.; width across eyes, 2.57 mm.; width across 
abdomen, 2.17 mm. 

Color. Body whitish green with ver>- little yellow. Head and thorax are 
generalh' of the same color as the body. 

Structural characteristics. Lateral angles of the first pair of wing pads 
are more produced and as long as the second pair. Second pair are also 
greath' enlarged both in width and length, reaching the third segment of the 
abdomen. Prothorax has the adult form with its emarginate posterior margin. 
Median portion of mesothorax, is produced caudad until it almost touches the 
posterior margin of the metathorax. Metathorax also slightly i)roduced caudad. 
On the metathoracic leg indication of the third tarsal segment is shown; the 
two rows of spines on the distal end of the tibia are also present, and the rows 
of spines on the distal end of each segment of the tarsus are faintly visible. 
The two front pairs of legs still have two segments in the tarsi and have two 
rows of spines on the distal end of the tibiae which are not present in the adult. 

NOTES ON THE DIFFERENT STAGES. 

Structural differences between the first and second instars were 
hard to discover. The chief difference is in the size, although a few 
minor characters are sometimes evident. In the second instar the 
apex of the labium just comes to the first coxa, but in the first 
instar it extends to the second coxa, or at least between the coxae 



538 The University Science Bulletin. 

of the first pair of legs. In the second instar the length of the 
mesothorax and metathorax together is approximately equal to 
that of the prothorax, while in the first stage the prothorax is longer 
than the other two together and the mesothorax is slightly longer 
than the metathorax. 

From the second to fifth instars color variation in the head and 
thorax can be found. Some specimens in each instar were both 
collected and reared in which the head and thorax were of the same 
color as the body. Others had head and thorax both washed in 
dark reddish brown, while still others had a pale prothorax with a 
dark mesothorax and metathorax, so that the thorax appeared to be 
distinctly banded. In the third and fourth instars the bands were 
of such dark-brown pigment that they appeared as black bands to 
the naked eye and made the nymphs very conspicuous. Apparently 
this color variation has no relation to sex, since both sexes have 
been found to have these different combinations. 

On the fourth, fifth and sixth abdominal segments of the first 
instar appear two large, oval, orange spots. These spots make the 
first-instar nymphs more conspicuous and more vividly colored 
than any of the other instars. They probably indicate the loeation 
of the spittle glands which secrete part at least of the viscid fluid 
of the foam. Garman (1921) says that they appear as yellow spots 
on the side of the abdomen of Philcenus, and Osborne (1916) found 
them as black spots in another species. Both writers state that the 
glands are found on the seventh and eighth segments, but in 
Lepyronia they are found on the above-mentioned segments. On 
the other hand, Kershaw (1914) says that the fluid is produced by 
the anterior or smooth portion of the malpighian tubes. The whole 
question is an interesting matter for further investigation. 

ECONOMIC IMPORTANCE. 

In general froghoppers are not considered economically important, 
since they are found feeding chiefly on weeds and wild plants. 
There are a few, however, which have been reported as doing con-' 
siderable damage to certain cultivated host plants. Osborne (1916) 
thinks that the damage caused by froghoppers has been over- 
looked. According to him, Philcenus spiwiarius L. considerably 
affects clover, for he noticed that the plants which are attacked 
wither and do not bear heads. He also has observed Philcenus lin- 
eatus L. being so abundant on grass and hay crops as to occasion 
distinct losses, timothy and redtop being most frequently injured. 



Doering: Lepyronia quadrangularis. 539 

Ashley (1919) reports an Aphrophora in England which causes 
serious damage to roses during June and July by weakening the 
young shoots and buds by extracting the sap. Harris (1862) re- 
ports Clastoptera protons as doing great injury to the cranberry 
crop in some parts of Massachusetts. The most destructive frog- 
hopper is the sugar-cane froghopper, Tomaspis varia Fabr., which 
is estimated by Urich (1913) as destroying ten per cent of the sugar 
crop of Trinidad. The damage which is done is called "blight" and 
consists of a withering of the leaves of the canes and a stunting of 
the stem. This is accomplished chiefly by the nymphs feeding on 
the roots, although the feeding of the adults on the leaves of very 
young plants may also cause it. Williams (1923) reports Clastop- 
tera theohromce Wills as destroying the flowers of the cacao in 
Panama. 

No records of serious damage caused by Lepyronia quadrangularis 
have been found. Fitch (1856) records it on grape. The writer has 
found it on several economic plants, such as timothy, black rasp- 
berry, wheat and sweet-clover. Most of these, excepting wheat, 
were isolated plants or small groups of them, and therefore were 
not of much importance. In the case of the wheat, however, the 
matter was more serious. A good many stalks of wheat were 
found bearing spittle masses. These occurred chiefly at one side of 
the field next to the weeds, bordering a woods, and evidently they 
had migrated from the weeds to the wheat. In all cases where the 
nymphs were feeding on the wheat the heads were half the size of 
the good heads. Nine good heads were averaged, and the number 
of kernels was thirty-four to a head. Eleven poor heads were 
averaged, and the number was only ten, which makes an ap- 
proximate loss of 33 per cent. From these observations, as well as 
those on the life history, the following conclusions can be drawn: 
That the insects by sucking the plant juices really cause consider- 
able damage to the host plant; and they may become a menace 
to certain cultivated crops by their ability to migrate from one plant 
to another, and because, unless disturbed, they are so free from 
parasites and predators,- they may greatly increase in number. 



540 The University Science Bulletin. 

EXTERNAL ANATOMY. 

Apparently little work has been done on morphology within the 

family Cercopidae. In no case throughout the literature available 

for study has a thorough investigation of the structure of the entire 

body been made. One paper on the alimentary canal of a cercopid 

by Kershaw (1914), one on the respiratory system of a nymph of 

the Aphrophorinae by Sulc (1910) , and one by Metcalf (1916) on the 

wing venation of Cercopidae, are the most extensive works to be 

found. Other brief references to the morphology of the Cercopidae 

were found in a number of articles dealing with the homologies of 

certain special parts of the body among the families of Homoptera, 

namely, Taylor (1918) on the thorax, Kershaw and Muir (1922) on 

the genitalia of both sexes, Crampton (1922) on the male genitalia, 

and Hansen (1890) on the appendages and spiracles. One other 

article on the structure of Aphrophora spumaria by J. 0. Harper 

(Science Gossip, 1874) concludes the list, but is of little value 

morphologically. 

THE HEAD. 

The terminology used in the discussion of the head is based mainly 
on that of Comstock and Kochi (1902) , Martlatt (1896) , and Funk- 
houser (1917). 

The deposition of chitin is fairly heavy and uniform throughout 
the exoskeleton of the head. In fact, both dorsal and ventral 
surfaces are hard and brittle making dissection very difficult with- 
out first boiling in KOH. 

A dense prostrate pubescence, consisting of thin, grayish hairs, 
covers the entire surface of the head. This gives the appearance of 
a grayish bloom, which partly conceals the coloration of the insect. 

The head of Lepyronia quadrangular is (Say) resembles the other 
Homoptera in most of the essential parts, although it resembles some 
more closely than others. Perhaps it is more nearly like the Cicad- 
idae and Cicadellidae when such points as the position of the head in 
relation to the body and the shape and comparative size of the 
sclerites are considered. The protruding, enlarged sclerite with its 
striated borders, commonly called the front, of Lepyronia instantly 
reminds one of the larger, homologous sclerite in the cicada. Like- 
wise the heads of both these insects lie in nearly the same plane as 
the body, which condition is seldom found in other Homoptera out- 
side of the Cicadellidae. In Lepyronia quadrangularis, however, a 
flattening of the head dorsoventrally has taken place, with the result 



Dokring: Lepyronia quadrangularis. 541 

that the vertex and the front lie entirely on the dorsal surface and 
the remaining sclerites on the ventral surface. Thus the anterior 
margins of the front and vertex (pi. T.V, fig. 1) have been pressed 
against the anterior margin of the clypcus, which together form 
the cephalic and lateral margins of the head. The head, as is shown 
in the figure, projects straight forward on a line with the body and 
the beak projects backward and lies between the coxse when at rest. 

In the literature the most commonly accepted homology of the 
sclerites of the head are as follows: On the ventral aspect the large 
protruding sclerite (pi. LVI, fig. 2) is called the front; the smaller 
sclerite attached to its posterior margin, the clypeus; and the 
smaller triangular sclerite fastened to the clypeus, the labrum. The 
entire dorsal portion of the head (pi. LVI, fig. 1) is usually desig- 
nated as the vertex, although it consists of two sclerites. The larger 
of these occupies most of the entire surface with the exception of the 
broad invagination in the cephalomedial region, into which a small 
subrectangular sclerite, the tylus, is inserted. There seems to have 
been no attempt to homologize or account for the existence of the 
tylus, although it appears as a definite sclerite in the adult and is 
partially indicated in the later nymphal stages. 

Partly because of the lack of explanation with regard to the tylus 
and partly because of the suggestion of other workers in other 
groups, an investigation with regard to a different homology of 
parts was attempted. Funkhouser (1917) states that the clypeus in 
the Membra cidae is determined by the location of the anterior 
arms of the tentorium, which fully accords with the view of Com- 
stock and Kochi (1902), that the anterior arms of the tentorium 
arise as invaginations of the body wall between the clypeus and 
front. He also refers to Bentley's work (1900) on the Cicada, which 
was unjuiblished, wherein the latter shows that the large, protruding 
sclerite known as the frons is really the clypeus. Several other 
articles in support of this theory have been found. Marlatt 
(1896) calls the sclerite in question (pi. LVI, fig. 2) the clypeus; 
likewise Smith (1892). Bugnion and Popoff (1911) state:" "One 
of the characteristics of the clypeus is that it serves for insertion, 
with its deep face, of the anterior bundles of the dilator of the 
pharj^nx. A clypeus cleared in balsam shows in all Hemiptera the 
linear insertion of the striated membrane (of the pharynx) and the 
two sides of transverse lines which correspond to the insertion of 
nauscles. Among some species (Cicada) the transverse lines are 
indeed seen from the exterior and give a particular appearance to 



542 The University Science Bulletin. 

the sclerite. Without a doubt the clypeus of which they speak is 
the frons of other writers. 

According to these writers, then, a different nomenclature from 
the commonly accepted one for the sclerites of the head of the 
Cicada has been used. The same thing holds true for Lepyronia 
quadrangularis. The sclerites on the ventral surface of head would 
be labeled clypeus, labrum and epipharynx (pi. LVI, fig. 2). In 
figure 1, the vertex remains the same, but the tylus would be termed 
the front. 

A study of the tentorium and the position of the ocelli bear out 
this homology for both the Cicada and Lepyronia. As was men- 
tioned above, Comstock and Kochi (1902) showed that the ends 
of the suture between the front and clypeus extend to the invagina- 
tion which forms the anterior arms of the tentorium. From this 
it follows necessarily that the position of the tentorium would un- 
doubtedly prove the identity of -the front and clypeus in these in- 
sects. The tentorium (pi. LVI, fig. 3) is of practically the same 
shape in the Cicada and in Lepyronia. It consists of a slender 
transverse bar, which is the basal part or body of the tentorium, 
from which two long, slender, chitinous arms extend cephalad, and 
two short, posterior arms extend laterad. In the Cicada the basal 
portion is fairly narrow, not extending over one-third of the width 
of the occipital foramen; it occupies a central position in the head 
cavity. The anterior arms are by far the longest part of the 
tentorium and can easily be seen to run almost directly cephalad 
and slightly laterad to the point' which marks the cephalolateral 
angle of the large protruding sclerite or clypeus. The tentorium 
in Lepyronia quadrangularis varies somewhat from that of the 
Cicada, but is essentially the same. In the cercopid the basal part 
of the tentorium is more extensive, in that it occupies about two- 
thirds of the width of the occipital foramen. The anterior arms 
extend in much the same direction as the Cicada, but their extreme 
anterior portions have become somewhat modified, due, no doubt, 
to the peculiar compression of the head. At this point it would 
probably be clearer to follow the figure (pi. LVI, fig. 3). In this 
figure the head is in the exact position as in figure 1, with the ex- 
ception that the vertex has been removed in order to show the 
tentorium. The vertex appears to be very loosely attached to the 
tylus, for it can readily be lifted away. After the vertex has been 
removed the connection of the tylus and clypeus can easily be seen. 
The anterolateral corners of the clypeus are reflexed and the cephalic 



Doerixg: Lepyronia quadrangularis. 543 

margin of the rcflexed corner has become strongly fused to the 
lateral margins of the tylus. It is not strange that with the peculiar 
arrangement of these sclerites, as well as the compression of the 
head on the anterior portion, that the tentorium has assumed a 
peculiar position also. In the first place, the anterior portion of the 
forearm of the tentorium has become forked. The prongs are 
blunt at the end and are of unequal length. The sm,aller one is at- 
tached to the skeleton of the head at a point about midway on the 
clypeus (fig. 3) and just back of the antennal pit. If the tentorium 
stopped here it would indeed be hard to identify the clypeus, since 
it is equally as far from either of the anterior margins of the two 
sclerites in question. Even in that case it would seem more plausible 
that it should have migrated backward due' to the flattening of the 
head, already described, than that it should have migrated so far 
forward for no accountable reason. Fortunately, however, the other 
branch of the fork extends farther cephalad. It curves slightly 
mesad, following the free margin of the reflexed clypeus, although 
not touching it. In fact, it extends as far forward as the corners of 
the clypeus at the point where the latter is attached to the latero- 
posterior angle of the tylus. The tip of the branch appears to be 
attached by a membrane to the ental surface of the vertex. The 
significance of the position of the tentorium at this place in the 
attempt to, locate, the front and clypeus seems to be this: If the 
reflexed corners of the clypeus were bent back in their normal 
position and the front and vertex were laid out on the plane of the 
rest of the head sclerites, then the anterior arm of the tentorium 
would extend to the laterocephalic margin of the sclerite labeled 
clypeus in figure 3. The sclerites of the head from this view could 
easily be designated in succession as the vertex, front, clypeus, 
labrum and epipharynx without any intervening unexplained tylus. 
Another argmiient for such nomenclature is the position of the 
median ocellus and its relation to the frons. Referring again to 
Comstock and Kochi (1902), we find a statement to the effect that 
the frons, in the more generalized insects at least, bears the median 
ocellus. Funkhouser (1917) is of the opinion that in the Mem- 
bracidae the frons has disappeared and with it the median ocellus 
which it contained. While Crawford (1914), in his work with 
Psyllidae, states that in all cases where the frons is present it bears 
the anterior ocellus at its base or at the end nearest the vertex. 
In the Cicadidae the median ocellus is distinctly located on the 
dorsal surface of the head in the sclerite just above that which is 



544 The University Science Bulletin. 

generally called the frons. In Lepyronia quadrangidaris there is 
of course no median ocellus present, but if it were present it seems 
highly improbable that it would be located on the ventral aspect 
of the head, so far removed from the other two ocelli. Its position 
would be more likely the normal one at about the place where the 
tylus is located. 

The compound eyes (pi. LVI, fig. 1) are large but not very dis- 
tinct on the dorsal aspect, due to the fact that their dorsal surfaces 
are level with the vertex, and the color of both is somewhat the 
same. They are oblong in shape, broader at the anterior end and 
narrower towards the posterior. They are located in the extreme 
caudolateral angles of the head. 

There are two ocelli (pi. LVI, fig. 1) present, located about mid- 
way between the front and the pronotum and about as far from each 
other as from the compound eyes. 

The antenncB (pi. LV, fig. 2) are located on the ventral surface of 
the head in a hollow pit halfway between the eyes and the clypeus 
and slightly cephalad of the eyes. In general appearance they are 
small and inconspicuous, appearing to consist of a very short basal 
stump and a long, thin, hairlike part which extends some distance 
beyond the sides of the head. From specimens stained with eosin 
and mounted on slides, more details of structure could be observed. 
The basal segments, called collectively the shaft or peduncle, are 
three in number. The first of these is subcylindrical, about as wide 
as long, and is apparently the widest of the three. The second 
segment is likewise subcylindrical, but approximately twice as long 
as wide, with its basal end slightly narrowed and its broader distal 
end cup-shaped. The last of the segments of the peduncle is the 
shortest, oviform in shape, and fits into the concave end of the 
second segment by means of a little stalk. The jiagellum or ivhip 
consists of many segments, the exact number of which it was diffi- 
cult to determine. After the first twenty-nine segments, extending 
two-thirds of the entire length, the segmentation becomes indistinct. 
In general these segments are cylindrical, but vary somewhat in 
shape, being longer than wide toward the tip of the filament and 
about equal in length and width at the base. The sense organs of 
the antennae are located at the base of the filament and at the tip of 
the third segment of the peduncle. They consist of a small group of 
sensory pits, averaging about eight in number, and generally located 
close to the apex and somewhat to one side of the segment. Appear- 
ing to rise from the tip of this segment are three short, conspicu- 
ous thick spines, which are also sensory in function. 



Doering: Lkpyronia quaurangulakis. 545 

The vertex (pi. LVI. fig. 1), as has already been mentioned, 
makes up the largest area of the dorsal part of Ihe head, trans- 
versely oiH'upying all that part of the head between the eompound 
eyes and longitudinally all the part between the occiput and tylus. 
In length and width it is about equal. In the anterior portion of 
the vertex is the subquadrate invagination into which the front is 
inserted. The suture between the front and vertex is very distinct 
and lii'in. hut the parts of the vertex bounding the front laterally 
do not appeiir to be fastened to the latter, but only to fit down over 
the line of union of the front and the reflexed corners of the clypeus. 
The posterior margin of the vertex is broadly emarginate, into which 
the rounded margin of the prothorax fits snugly, while the lateral 
margins round to a blunt apex. 

The occiput (pi. LVI, fig. 1) is not visible until after the head has 
been removed from the body, since it occupies the central portion 
of the caudal surface of the head, or that part which lies against 
the cephalic surface of the pronotum. It is a single sclerite, forming 
the upper boundary of the occipital foramen and the posterior 
boundary of the vertex. Laterad the lower ends of the occiput fuse 
with the postgenae, so that the suture appears to stop at a point in 
a line with the inner margin of the eye. 

The postgence (pi. LVI, fig. 1) occupy the remainder of the caudal 
surface of the head. They are triangular in shape, with the sharply 
tapering apex extending between the vertex and occiput. The upper 
boundary of each postgena is formed partly by the lower margin of 
the eye and partly by that of the vertex. Its outer margin is the 
outer limit of the head and its lower end is fused with the occiput. 

The front;, or front (plate LVI, fig. 1), commonly spoken of as 
the tylus. as will be noted from the foregoing, is located on the 
dorsal surface of the head. It is subquadrangular in shape, about 
one-third the width of the vertex and about one-half its length. 

The clypeus (pi. LVI, fig. 2) is located on the ventral surface of 
the head. Its position has been determined by the fact that the 
forearms of the tentorium extend to its laterocephalic angles. 

The clypeus is enormously enlarged and protruding, subquad- 
rangular in shape, widest in the middle and narrower toward each 
end. In size it extends practically two-thirds the length of the head 
and over one-half its width. On each lateral margin is a row of 
parallel grooves separated from each other by a plain, wide, chitin- 
ous band. These are lighter than the surrounding parts and there- 
fore stand out clearly. The peculiar reflexed condition of the 



546 The University Science Bulletin. 

anterior corners and their connection to the front have already been 
described under the discussion of the tentorium. 

The labruni (pi. LVI, fig. 2) is a flask-shaped sclerite with its 
broad end attached to the clypeus and its apex truncated. In 
length it is half the size of the clypeus and in width equal to the 
width of the clypeus in the neighborhood of the clypeal suture, but 
narrows down to half the width at its distal end. The suture be- 
tween the clypeus and the labrum is indistinct along the median 
line, but is plainly indicated at the sides. The labrum is heavily 
chitinized and appears to be quite rigid. 

Attached to the caudal end of the labrum and lying on top of the 
base of the stylets is a small triangular sclerite. This is the epi- 
pharynx (pi. LVI, fig. 2). It is very thin and lightly chitinized, for 
which reason it is not readily seen. 

The gence (pi. LVI, fig. 2 1 occupy most of the lateral parts on 
the ventral aspect of the head. They are irregular in shape with 
their lateral margins following the contour of the compound eyes 
and their median margin touching the clypeus and the maxillary 
sclerite. Caudad they form the outer margin or boundary of the 
head, which also is the suture between the gena? and postgena?. 

The occipital foramen (pi. LVI, fig. 1) is the opening visible 
from the caudal aspect of the head. It is bounded dorsad by the 
occiput and laterally by the postgenai and mandibular sclerites. 

The rostrum (pi. LVI, figs. 1 and 2), or beak, is composed of the 
labium and two pair of stylets, which are modified maxillae and 
mandibles. The labium is composed of three segments. The basal 
segment is the shortest of the three and cylindrical in shape. The 
second segment is the longest, being about three times the length 
of the first and twice the length of the distal segment. The last 
segment is rounded at the tip, its margins seemingly more com- 
pletely folding around the stylets. The beak is straight and ex- 
tends backward between the middle coxa? for half their length. The 
labium is connected by a membrane to both the head and sternum 
of the prothorax, its point of attachment being just above the tip 
of the labium. The attachment to the head seems to be the weaker 
of the two, for when the prothorax is detached from the head the 
labium usually comes off with it. There is a groove along the ven- 
tral surface of the labium, in which repose the maxillae and man- 
dibles. 

The mandibles and maxillm, as in other Homoptera, are setiform 
or bristlelike. The mandible is connected to the head by a slender, 



DOERINU: I.Kl'VHOMA QIADRANGULARIS. 547 

chirinous i-od at a point wIum-c the caudal oorn(M's of postgense are 
joined to tlie mandibular sclerite. Tiie maxilla is fastened to the 
anterior rorncr of the maxillary sclerite, which places it cephalad 
of the mandible. Funkhouser (1917) states that in the Membracidse 
they are attached to the vertex, which differs from the condition 
found in most other insects, where they are attached to the post- 
genae. In Lcpyronia quadrangulari.s neither of these conditions 
occurs, but the close approximation of the mandibular sclerite to the 
postgena may be significant. 

In order to study the structure of the tips of the mandible and 
maxilla it was necessary to study them under high magnification. 
Both stylets are cylindrical in shape, tapering toward a blunt point 
and with their hidden bases broadening into flat plates. The outer 
margin of the distal third of the mandible is finely notched, w^iile 
the outer apical part only of the maxilla is modified into five dis- 
tinct, rounded teeth (pi. LVI, fig. 4). The maxilla? appear to be 
equal to the labium in length, but the mandibles are slightly longer, 
so that their tips protrude a short distance beyond the tip of the 
labium. 

The four stylets are closely associated, converging at the place 
where the labium is attached to the head. Here the mandibles be- 
come joined together by their inner margins, forming a shallow 
sheath in which the maxillae lie. The maxillae are also united, prob- 
ably forming a tube, as in the Cicada (Snodgrass, 1921), through 
which the sap is sucked up into the pharynx. The maxillae of 
Lepyronia, however, do not appear to be so firmly joined as in the 
Cicada, for their tips often spread apart, until from a superficial 
view, they appear to be on the outside of the mandibles. 

The maxillary sclerites (pi. LVI, fig. 2) are the large semicircular 
sclerites on the ventral aspect of the head, bounded laterad by the 
genae, caudad by the mandibular sclerites, and mesad by the clypeus 
and labrum. The cephalic tip' of each maxillary sclerite is con- 
nected to the maxilla by a slender, chitinized branch of the latter. 
These sclerites are homologous to the lorae of the Cicadellidae. Snod- 
grass (1921) considers them as definite sclerites, but attaches no 
name to them. 

The mandibular sclerite (pi. LVI, figs. 1, 2) extends from the 
caudal angle of the genae and postgenae to the base of the labium. 
Each sclerite is elongate, narrow at the base, but widening in the 
middle to a flangelike projection and tapering to a tonguelike apex 
which partially folds around the base of the stylets. The interior 



548 The University Science Bulletin. 

surface of this flangelike part forms a pocket or fold, called by 
Martlatt (1895 » a pseudo-sheath, which receives and partially pro- 
tects the mandible. It is connected to the mandible, as in the 
maxilla, by a chitinous arm of the stylet. It may be possible that 
the mandibular sclerite is comparable to the sclerite found in lower 
orders of insects, which is termed the trochantin of the ynandible by 
Comstock and Kochi (1902) or the basi'mandibiihe by Crampton 
(1921). 

THE THORAX. 

The prothorax, as is shown in (pi. LVI, fig. 6) is weakly attached 
to the mesothorax, but is closely associated with the head, since it 
invariably comes off with the head when the latter is removed from 
the body. The mesothorax and metathorax, however, are firmly 
joined and the sclerites somewhat overlap on the dorsal surface, 
making identification of the sclerites rather difficult. 

There is no evidence of cervical intersegmentalia in Lepyronia, or 
of any structures which might suggest them. The cephalic surface 
of the prothorax appears to fit directly against the caudal surface of 
the head without any intervening structures which might be termed 
these sclerites. 

THE PROTHORAX. 

The notum (pi. LVII, fig. 1) of the prothorax is a large, fiat 
sclerite occupying the entire dorsal surface of this segment. It is 
twice as long as wide and has its short, lateral margins nearly paral- 
lel. The anterior margin is broadly rounded. The posterior margin 
is deeply emarginate on the produced middle third. The edges of the 
notum are greatly deflexed so that they unite with the pleuron on 
the ventral surface. 

From a cephalic aspect of the pronotum (pi. LVII, fig. 5) can be 
seen a narrow sclerite, which probably corresponds to the sclerite in 
the Cicada termed by Taylor (1918) the pretergite. Mesad this 
sclerite is very narrow, but laterally it widens out to four or five 
times its median width. It lies at right angles to the notum and is 
connected to the pleuron by a tapering projection. The pretergite 
is not as heavily chitinized as the notum and pleuron. 

The pleuron (pi. LVII, fig. 3) of the prothorax is closely joined 
to the deflexed part of the notum without a distinct line of connec- 
tion. It is divided longitudinally by a very indistinct pleural suture 
into two sclerites, the epimeron and episternum. The episternum, 
the anterior of the two, is roughly rectangular in shape and about 
twice as long as wide. The epimeron is slightly larger than the 



DoEHiNt;: Lki'Ykoxia QrAi)HA.\(irLAKis. 549 

cpistcrnum. more irrciiuhu' in shape and extends dorsad into a 
tajiei'ing point. The episternuni is connected to the sternum by a 
naiTow prcco.ralr bridge (pL LVII, fig. 4), called the precoxale by 
Taylor (IDlSi. It is fused to the ventroentad surface of the epi- 
sternuni and it is not distinctly separated from the sternum. The 
epimeron is connected in much the same manner by a postcoxale 
bridge (pi. LVII, fig. 4) which, however, runs to the laterodorsal 
angle of the epimeron instead of the ventral. 

The sternum (pi. LVII, figs. 4 and 5) of the prothorax is small 
and about equal in length and width. It consists of a single sclerite, 
which is folded and curved in such a manner that it is difficult to 
describe its exact shape. 

The anterior margin of the sternum is bent back against the 
caudal surface of the sclerite (pi. LVII, fig. 4), so that in an un- 
cleared specimen it appears as a chitinous ridge or distinct sclerite. 
In the drawing of the cephalic aspect this has been straightened out 
into a natural position. The lateral margins of the sternum (pi. 
LMI, fig. 5l curve cephalad, thus forming a semicylindrical cavity 
into which the membrane connecting the labium with the sternum is 
fastened. During the process of sucking the sap the labium is 
probably drawn up into this trough in order to not interfere with 
the action of the stylets. The anterolateral angles of the sternum 
bear the furca (Taylor, 1918), which are the processes for the at- 
tachment of muscles. 

The trochantin (pi. LVII, fig. 5) is a small but very distinct knob- 
like sclerite just below the ventral end of the episternum. 

THE MESOTHOR.\X. 

The mesothorax (pi. LVII, figs. 7 and 8) is closely connected with 
the metathorax, the latter being partially covered both on its dorsal 
and ventral surfaces by overlapping parts of the mesothorax. The 
mesothorax, likewise, is covered by the pronotum with the exception 
of the scutellum, the anterior margin of which follows the deeply 
incised posterior margin of the pronotum. The membranous ster- 
nellum of the mesothorax is fused with the presternum of the 
metathorax so that it is difficult to differentiate the two. 

The mesonotum (pi. LVII, fig. 7) is divided into four distinct 
areas or sclerites. The first of these is the prescutum, which occu- 
pies the anteromesal portion and is bounded laterad by two longi- 
tudinal sutures which curve mesad for a short distance, thus 
indicating the caudal boundary of the sclerite. The prescutum 
is heavily chitinized in spite of the fact that it is entirely covered 



550 The University Science Bulletin. 

by the pronotum. On its anterior margin it bears the anterior 
phragma (pi. LVII, fig. 7), a semimembranous, bilobed, narrow 
structm-e which is connected to the pronotmn by a membrane. 

The scutum (pL LVII, fig. 7) of the mesothorax is an irregular 
sclerite, occupying the entire lateral regions of the notum. It ap- 
pears to consist of two sclerites, since the anterior point of the 
scutellum almost reaches the caudal boundary of the prescutum. 
At its laterocaudal angles the scutum is deeply incised to allow for 
the wing processes. 

The scutellum (pi. LVII, fig. 7) is composed of a large shield- 
shaped piece, which is visible externally and two narrow lateral 
portions which connect with the anal margins of the wing and are 
not visible externally. The large median part is separated from 
each lateral part by a narrow groove into which the deflexed claval 
portion of the wing fits. 

There are several small sclerites to be found connecting the base 
of the wing with that of the notum. On the anterolateral corner of 
the scutum is a small piece, called by Taylor (1918) the suralare. 
It is marked by a cleft which extends forward. As in the Cicada, 
another cleft extending backward marks off a similar piece. Just in 
front of the incision made by these clefts is a small triangular, free 
plate, the notopterale. Caudad of this plate, imbedded in the mem- 
brane, is a larger, subrectangular plate called the adanal pterale. 
There is no indication of the presence of the tegula, which agrees 
with the conclusion of Taylor that the tegula is not present in the 
families of Homoptera outside of the Fulgoridae and Cicadidse. 

Cephalad of the wing the notum is connected to the pleuron by a 
narrow prealare bridge (pi. LVII, fig. 7). The postscutellum is con- 
nected to the epimeron by a similar postalare bridge (pi. LVII, 
fig. 6). 

The postscutellum is a narrow sclerite, entirely hidden by the 
scutellum and is membranous in structure. 

The pleuron (pi. LVII, fig. 6) is somewhat more complicated than 
that of the prothorax. The pleuron, in fact, forms a major part of 
the ventral body wall, so that the shape of the episternum and epi- 
meron can best be seen from a ventral view (pi. LVII, fig. 10.) The 
pleural suture (pi. LVII, fig. 6), is ciuite distinct, consisting of an 
almost straight line extending from the base of the prealare bridge 
to the base of the coxae. 

The epistermnn (pi. LVII, fig. 10) is a single sclerite, which is not 
divided into an anepisternum and katepisternum unless a diagonal 
elevated ridge is an indication of this division. The episternum 



Doering: Lepyronia quadrangularis. 551 

is irregular in shape, about as wide as long;, and approximately 
equal to the lengtli of the sternum. It is separated from the sternum 
by a distinct transverse suture. There is no precoxale bridge. 

The epimeron (pi. LVH, fig. 10) is an elongate selerite extending 
the entire length of the mesothorax. It is widest just behind the 
middle when \ie\ved ventrally, and bears the postalare bridge, 
connecting it with the postscutellum just cephahid of its broadest 
point. The epimeron is partly divided longitudinally by a suture 
which runs parallel to the pleural suture, but which extends for only 
lialf its length. Like the episternum, it is fused to the sternum, 
being separated by the pleural suture. 

The sternum (pi. LVII, fig. 10) of the mesothorax is composed of 
three sclerites — the presternum, the sternum proper, and the 
sternellum. The anterior one is the presternuni (pi. LVII, fig. 10), 
which is a narrow collarlike selerite, bounded laterad by the 
episterna and caudad by the sternum itself. It is slightly less 
chitinized than the sternum, especially towards its lateral margins. 
Along its anteromedian margin is a broad indentation into which 
the troughlike sternum of the prothorax is inserted. 

The sternum (pi. LVII, fig. 10) is the major selerite of the ventral 
part of the mesothorax. It is a bilobed selerite, each lobe being 
somewhat inflated. They are separated from each other by a heavy, 
dark median line which resembles a suture. Just before this line 
reaches the caudal margin of the sternum it extends laterad as a 
short cleft which marks off a little flap on each lobe. The sternum 
is heavily chitinized and very dark colored. It is closely associated 
with the pleural sclerites, as was mentioned above, but is separated 
from both by very distinct and heavy sutures. 

The sternellum (pi. LVII, fig. 10) is a partly membranous, partly 
chitinous selerite, lying directly caudad of the sternum. It is a 
rather ill-defined region, somewhat semicircular, with its caudal, 
convex margin joined to the concave cephalic margin of the pre- 
sternum of the metathorax. The chitin is deposited along the 
middle line of this selerite as a broad, longitudinal band which sends 
out two lateral arms at its cephalic end which are entirely hidden 
by the overlapping sternum, and two lateral arms at its caudal 
end. The part of the sternellum which is chitinized is not very 
heavy, due to the fact that the large coxae entirely cover it. 

The trochantin (pi. LVII, fig. 10) is a very small but distinct 
selerite, located between the laterocaudal angle of the sternum and 
the laterocephalic angle of the coxa. 



552 The University Science Bulletin. 

the metathorax. 

Tlie connection of the metathorax to the mesothorax has already 
been noted. It is weakly joined to the abdomen, not only because 
it is connected to the abdomen by a membrane, but also because 
the first segment of the latter is partly membranous itself. It is 
similar to the mesothorax in general make-up, especially in the 
number of sclerites. 

The nohmi (pi. LVII, fig. 91, as in the mesothorax, is composed 
of four sclerites — the prescutum, scutum, scutellmn, and post- 
scutellum. The prescutum is peculiar in that it is entirely internal, 
extending directly into the body cavity and lying at right angles 
to the scutum. It is a very large, bilobed sclerite, heavily chitinized, 
and is used for attachment of large bundles of muscles. 

The scutum is the large, heavily chitinized sclerite found on the 
dorsal surface of this segment. It is rectangular in shape and its 
width is one and one-half times its length. Along its median line 
is a heavy, black line which is the external indication of an infolding 
of chitin, which also serves for muscle attachment. 

The scutellum of the metathorax is much reduced, consisting of 
a small, median portion, which is produced laterad into a narrow 
band. The latter is directly connected with the anal margin of the 
wing. 

The postsciitcllum is a flat chitinous sclerite, which is extremely 
narrow mesad. but which expands laterad into two tapering side 
processes. The cephalic margin of the postscutellum is attached 
to the anterior margin of the scutellum, which is the suture sep- 
arating the scutum from the scutellum. The diagonal, lateral angle 
of the postscutellum is directly attached to the margin of the 
cpimeron and no postalare is present, as in the Cicada. 

The pleuron (pi. LVII, fig. 8) is very well developed, being by 
far the largest part of the metathorax. It not only occupies all the 
pleural regions, but extends into the dorsal and ventral regions and 
forms a major part of the metathorax. 

A distinct pleural suture extends the entire length of the pleuron. 

The episternuin is an undivided sclerite, although on its caudal 
end is the beginning of a suture which runs parallel to the pleural 
suture. The episternum is the largest of the pleural sclerites, 
occupying the ventral half of the pleuron, the lateral third of the 
venter, and the laterocephalic angle of the dorsum, where it curves 
around the wing process. 



Doerixg: Lki'yronia quadrangularis. 553 

The cpimcron is likewise a large sclerite, but it is found chiefly 
on the lat(M-al and dorsal parts of the segment. It occupies the 
dorsal half ol' the pleuron and lateral part of the dorsum, where 
mesad it touclu^s the lateral margins of the scutum and postscutel- 
luni (J)!. lA'II. fig. l)f. The lateral extensions of the metascutellum, 
which arc joined to the axillary coi'd of the wing, lie on top of the 
epinu'i'on. From a lateral view (pi. LVII, fig. 8) the indentation 
made hy I he wing process can easily be seen. Ventrally the epi- 
meron shows as a })late forming the caudal boundary of the epister- 
num. A <uture divides the sclerite into an upper and lower portion. 
A fiaplike process is present in the lower epimeron of Lepyronia, 
which is probably similar to that found by Taylor (1918) in an 
Aphrophom. 

The stennnn. (pi. L^TI, fig. 11) is small when compared to the 
sternum of the mesothorax. The presternum of this segment is 
l)artly membranous, with a weak deposition of chitin in its median 
and caudal portions. It is roughly crescentic in shape. 

The sternum is entirely membranous and is divided into two dis- 
tinct halves by a very narrow, chitinous band, which appears to be 
a caudal extension of the chitinous part of the presternum. 

The sternelhtm appears to be a narrow, chitinous rod extending 
between the epimera, with which it appears to be fused. It is 
located beneath the posterior coxse, and for this reason is more 
easily seen from a dorsal view than from a ventral view (pi. LVI, 
fig. 7). 

The trochanfin is a knoblike structure terminating the caudal 
end of the pleuron. It is termed by Taylor (1918) the meron, or at 
least he assigns this term to a similar structure in an Aphrophora, 
but does not account for a trochantin. 

THE WINGS. 

The jorewing (pi. LVIII, fig. 4) of Lepyronia is thick, opaciue 
and rather tough. Its entire surface is broken up into fine reticula- 
tions and is covered by a grayish pubescence, similar to that of the 
head. These conditions, together with the even coloring of the 
wing, entirely obscure the venation in uncleared specimens. It was 
only in half-bleached specimens that the veins could be traced at 
all. The venation of both wings here studied was found to be 
almost identical with the venation of the same species as described 
by Metcalf fl9]7), 

Costa is a single, unbranched vein which forms the costal border. 



554 The University Science Bulletin. 

Subcosta is so closely related to radius, at least distally, that it 
only appears as a definite vein along the center of its course, where 
it is widely separated from radius and thus forms an elongate, oval 
cell. Radius has three branches, which are R^ and R, + 3 and R^ + .,. 
Media is unbranched and is closely connected with cubitus, at least 
at its base. Cubitus is two-branched. There are three anal veins 
present, the first one of which is united with cubitus 2 on the 
margin of the wing. 

In the hindwing the veins show up very plainly, being brown in 
color and quite thick, while the cells of the wing are membranous. 
Radius is two-branched, the branches being R^ + ^ and R^ + 5. Media 
is unbranched and cubitus is two-branched. Three anal veins are 
likewise present in this wing, the first one being closely connected 
with cubitus basally, \\4iile the third one is branched. 

Part of the anterior margin of the posterior wing is produced into 
a triangular projection whose outer margin bears from four to six 
stout spines or hooks. Also on the ental surface of the tegmina 
just in front of subcosta is a little elevated keel. According to 
Hansen (1890) these hooks and keel are complementary structures 
and contribute to the steadiness of the wings. 

THE LEGS. 

The three pairs of legs have a general similarity with respect to 
position on the body, development, relative size, and number of 
segments. When in action the front legs usually point forward and 
the two last pairs point backward I pi. LV, fig. 1), but when at rest 
they are drawn up under the sloping tegmina (pi. LV, fig. 4) so that 
they are invisible from above. The hind pair is the longest and 
the other two are about equal in length. All are well developed 
as a result of their jumping habit. The number of segments is the 
same in each, consisting of coxa, trochanter, femur, tibia and a 
three-segmented tarsus. They are blackish brown in color, espe- 
cially the proximal segments, but gradually grow lighter toward 
the distal ones. All three pairs are covered by a fine pubescence. 

The coxa (pi. LVIII, fig. 3) of the front leg is stout and thick in 
comparison with the other segments of the leg, but it is not as large 
as the coxae of the other two pairs. It is smooth and about equal 
in length and width. It is almost cylindrical in shape, although 
its proximal end broadens out transversely and thus gives the effect 
of a flattened plate. The two anterior coxse are separated from each 
other bv the width of the beak, which fits down between them. 



Doering: Lepyroxia quadrangularis. 555 

The trochanter is a small elbow-shaped piece whieh is attaehetl to 
the \-entromesal angle of the eoxa. It is slightly constricted at its 
cephalic end and is about one-fourth the size of the coxa. 

The femur is a cylindrical-shaped segment which is somewhat 
swollen at the proximal end, but gradually becomes narrow distad. 
It is the stoutest of tlii' leg segments outside of the coxa. The 
trochanter is attached to its niediocephalic angle by a diagonal line 
of connection. Its distal end bears a small groove into which the 
tibia is inserted, while the lateral margins of the groove expand 
into a platelike structure on cither side of the tibia for the purpose 
of strengthening the joint. 

The tibia resembles the femur in general shape. It is cylindrical, 
smooth, and approximately the same length as the femur, but its 
width is only half that of the femur. The proximal end, which 
fits into the grove of the femur, is elbowed. There are no spines on 
the front tibia. 

The tarsus is three-segmented. The first two segments are about 
equal in length, triangular in shape and larger ventrally tluin dor- 
sally. 

The tarsus is terminated by two equal, heavily chitinized claws 
(pi. LVIII, figs. 6 and 9). The claws are broad at the base and 
gradually taper to an obtuse blunt point. The pulvillus or cm- 
podium, accoi'ding to Hansen (1890), is very conspicuous and 
greatly extended. On its dorsal surface it bears a large, bilobed, 
chitinous plate, and just below this a bristle (pi. LVIII, fig. 6). 
On the ventral side (pi. LVIII, fig. 9) are two narrow, longitudinal 
bands which run parallel to the margins of the claws. These bands 
and part of the claws appear to be attached to the third segment of 
the tarsus by a lightly chitinized band or sttilk. 

The middle legs (pi. LVIII, fig. 2) are attached directly to the 
sternum by the broad, transverse margins of the coxa^. The trochan- 
tin of the mesothorax is much smaller than that of the prothorax and 
is situated on the extreme laterocephalic angle of the coxa just above 
the meracanthus. The median leg. in most respects, is similar to 
the fore leg, being almost identical in size and shape of segments 
with the exception of the coxae. The latter are somewhat larger 
than the anterior coxse and their median margins are almost con- 
tiguous. They have on their lateral margins a large, flattened 
process or meracanthus which is not found on the other four legs. 

The posterior coxce fpl. LVIII, fig. 8) are more broadly attached 
to the body than even the intermediate coxae are, since they extend 



556 The University Science Bulletin. 

across the entire width of the sternum. The sternellum and the first 
segment of the abdomen is completely hidden by them, and part of 
the second and third segments of the latter are also covered. They 
are contiguous along their inner margins, which fact, together with 
their broad, basal attachment, gives them the appearance of great 
rigidity. In shape they are roughly rectangular with their latero- 
cephalic angle somewhat extended. 

The trochanter of the hind leg is similar to those of the first two 
pairs of legs, in that it is an elbow-shaped segment and of the same 
size as the others. 

The femur is a smooth, cylindrical-shaped segment, diagonally 
attached to the trochanter. On its proximal lateroventral margin is 
an obliciue protuberance (pi. LVIII, fig. 7), which, according to 
Hansen (1890), occurs in all Cercopidae, but in no other Homoptera. 
The knee joint is strengthened, as in the two anterior pairs of legs, 
by the groove and side plates of the femur, which are very prominent 
and show plainly in figures 7 and 8 (pi. LVIII). 

The tibia of the hind leg is greatly lengthened. Basally it is c^uite 
narrow, but broadens to twice its width distad. Along its outer 
margin it bears two large, thick spurs, which are of taxonomic im- 
portance in distinguishing the family. The second of these is ap- 
proximately twice the size of the first in both length and thickness. 
The distal end of the tibia bears two rows of thick, sharply pointed 
spines. The average number of spines, after counting twenty speci- 
mens, was seven on the top row and eight on the lower. These spines 
are shiny black at the tip and from between each two of them arises 
a long, silken hair which is twice the length of the spines. 

The tarsus is composed of three segments, the first two of which 
are similar in shape to the tibia, being narrow at the base and 
spatulate at the tip. They also bear a row of spines on their distal 
ends, which are like those of the tibia in shape and color but which 
are only half as large. The average number of spines on the first 
segment is seven, and on the second, nine to eleven. The distal 
segment of the tarsus is swollen but not spatulate. 

THE ABDOMEN. 

The number of segments in both the male and female abdomens 
is eleven plus a telson. From a dorsal and lateral view the abdomen 
is not visible, since the sloping tegmina completely hide it. It can, 
of course, be easily seen from a ventral view, but owing to the en- 
larged hind legs, especially the coxae, and the fact that its segments 



Doering: Lepyroxia quadrangularis. 557 

are more or lo?s telescoped, few details of structure can be made out 
without first removing the legs. Its color is like that of the rest of 
the body, and is uniform throughout with the exception of the 
sutures, whicii are somewhat lighter in color, indicating their mem- 
branous nature. The abdomen is A-ery wide at its base, but tapers 
to a pointed apex, dui' to the fact that the segments decrease in 
size from front to rear. Each segment overlaps the following one, 
whicli produces a telescopic effect. The segments are divided into 
three general regions — tergum, pleuron, and sternum. The tergum 
occupies all the dorsal region, while the jileuron and sternum are to 
be found on the ventral surface. 

The ventral, dorsal and lateral views of the male abdomen are 
shown on plate LIX, figs. 1, 2 and 3. The first segment was diffi- 
cult to find, owing to its irregularity of shape and its half-mem- 
])ranous structure. The tergum is partly membranous and partly 
chitinized. although the chitin is very weak even where it is present. 
Along its cephalic margin is a transverse, chitinous band, which 
mesad is very narrow but which is broadly expanded laterad. This 
anterior band is separated from a similar posterior, chitinous band 
by a strip of membrane. The first tergum appears to be attached 
to the postscutellum by a membrane. 

The pleuron (pi. LIX, fig. 2) of the first segment is greatly re- 
duced and occupies a peculiar position on the abdomen. It is a 
small, triangular, heavily chitinized plate lying on the laterocaudal 
angles of the tergum. This is the only pleuron visible from the 
dorsal view of the abdomen, but is easily recognized as such by the 
distinct spiracle located near its upper margin. 

The sternum is also greatly modified. Its anterior margin is pro- 
duced into a flaplike structure which bends forward on the re- 
mainder of the segment, due to pressure from the thorax (dotted line, 
pi. LIX, fig. 1). The lateral parts of the first sternum are mem- 
branous and deeply depressed where the chitinous angles of the 
thorax fit down against it. 

The tergum of the second segment is entirely chitinous, not ex- 
tending across the entire dorsal surface of tlie alxlomen, but with 
its lateral, rounded margins extending cephalad and bounded by 
the cephalad-projecting corner of the thiixl tergum. 

The pleuron (pi. LIX, fig. 1) of the second segment is of the same 
shape as the pleuron of the first, but is somewhat smaller. The 
relation of the two can be seen from a lateral view (pi. LIX, fig. 3). 
The two bases of the triangles adjoin each other, while the apices 



558 The University Science Bulletin. 

extend dorsad and ventrad, respectively. The spiracle of this 
pleuron is difficult to locate, since it lies near the mesal margin of 
the pleuron and is visible only from a ventral view of the abdomen. 

The sternum of the second segment is an elongate, narrow, chiti- 
nous band which tapers laterad to a fine point. Superficially this is 
taken as the first sternite, since the latter, because of its membran- 
ous condition, usually is pulled off with the thorax when it is re- 
moved. 

The third to eighth segments are practically all alike, being ring- 
like, in form and differing only in size. The terga are heavily chiti- 
nized and occupy the entire dorsal surface. The pleura are roughly 
rectangular, somewhat inflated, and with both lateral and median 
angles slightly rounding. The pleura are wider than either the 
sterna or terga. The sterna are elongate, rectangular plates forming 
the real ventral wall of the abdomen. 

The pygojer or gononiere is the modified ninth tergite, which is a 
rounded, bulblike structure, bearing on its dorsal surface the anal 
tube. The ninth sternite forms the plate beneath the genital ap- 
paratus, and is called by Crampton (1922) the hypandrium (pi. 
LIX, fig. 1). These make the ninth segment a very prominent seg- 
ment of the abdomen. 

The anal tube (pi. LIX, fig. 2), as was stated above, is situated 
on the dorsal surface of the abdomen, apparently fitting into a cir- 
cular depression and connected to the pygofer by a membrane. It 
is composed of two segments, which represent the tenth and 
ele^'enth abdominal rings or uromeres. The tenth uromere is by far 
the larger and is termed by Crampton the proctiger. The eleventh 
segment bears the telson, which is triangular in shape and is divided 
in the middle into two distinct halves by the anus. 

The female abdomen (pi. LIX, figs. 4, 5, and 6) is similar to that 
of the male in most details. The first tergum is both membranous 
and chitinous, but differs from the male in that there is only a 
single, irregular band of chitin along its cephalic margin instead of 
two. The first sternum and pleuron are much the same as in the 
male. 

The second to seventh segments of the female show little difference 
from those of the male. The eighth sternum, however, has been 
slightly modified to admit the ovipositor. It is divided into two 
distinct halves by the bases of the ovipositor valves, which extend 
cephalad until they reach the caudal margin of the seventh sternum. 
Each plate of this sternum is an inverted right-angled triangle, 



Doerixg: Lepyroma qiadrangularis. 559 

with ils caudal margin sonirwhat cuvving. The eighth pk'ura and 
tergum appear quite nonual. 

Tlu^ ninth segment, or pinjojir, is greatly enlarged and intlaled, 
even more so than in the male. The anal tube is borne on its dorsal 
surface, but occupies a more caudal position than in the male. For 
this reason the pygofer is more bulblike, extending around on the 
ventral surface, and thus making up a large share of the posterior 
end of the female abdomen. Its median margins do not meet on the 
ventral surface, since the valves of the ovipositor extend between 
them. There appears to be no ninth pleuron externally, as its 
position is occupied by the tergum, but a remnant of the nintb 
sternum (pi. LIX, fig. 6) is to be seen, forming the two slender basal 
parts of the dorsal valves of the ovipositor. 

The anal tube, as in the male, consists of the tenth and eleventh 
segments plus the triangular telson, and is attached to the posterior 
region of the pygofer by a membrane which fits into a circular 
opening. The eleventh segment is greatly reduced, however, being 
about one-eighth as long as the same segment of the male. 

THE MALE GENITALIA. 

From an external view of the abdomen part of the so-called 
"iHternal genitalia," as well as the "external genitalia," are visible. 
The organs usually included as "internal genitalia" are the paired 
styles, the cedagus and the connective. The parts wdiich are exposed 
are the apical two-thirds of the styles. These are plainly visible on 
the dorsal surface of the abdomen and have no protective covering 
of any sort. 

The basal parts of the styles, the connective, and the oedagus are 
situated in a genital or terminal chamber formed by the pygofer 
and anal tube. Since this chamber is really not closed, but is open 
beneath the anal tube, these parts are not really internal, but are 
only designated as such for convenience. 

There is no indication of the presence of the diaphragm, which 
Gifford (1922) found to be present in the genital chamber of the 
Delphacidse. In fact, the entire genital chamber of these insects 
varies considerably from that of Lepyronia. 

The genital plates (pi. LIX, fig. 1) are two valvelike processes 
on the ventral surface of the abdomen. They have been given vari- 
ous names, such as hypovalvae, ventral plates, and genital plates. 
They are large, prominent flat plates, occupying most of the apical 
end of the abdomen, and arc broad at the base but taper to a pointed 



560 The University Science Bulletin. 

apex. These plates, as is shown from nymphal development, arise 
from a genital area on the ninth segment. They are indistinctly 
fused with the caudoventral margin of the pygofer or with the area 
termed by Crampton (1922) the hypandrium. The lateral margins 
of the hypandrimn are indistinguishably united with the lateral 
margins of the pygofer. The plates are separated along their mesal 
lines by a long cleft which extends nearly to the eighth sternite. 

The styles (pi. LXI, figs. 7 and 8) have been given various names, 
such as gonistyli, claspers, or genital styli. They are very irregular 
in shape. At their cephalic end they taper to a very slender point, 
while their caudal end is truncate. The widest part of the style is 
at the point of attachment to the connective. Just cephalad and 
caudad of this point it is constricted, but it broadens out again 
caudad of the latter constriction into a second enlarged part. At 
this particular point on the dorsal surface of each clasper arises a 
prominent hook, the tip of which extends in a cephalomesal direc- 
tion. From a lateral view of the genitalia (pi. LXI, fig. 5) the 
nature of the relation of this hook to the style can best be seen. 
Just distad of each hook is a slight notch in the clasper. The apex 
of the organ bends mesad in a distinct cur^•e. The styli are con- 
nected to the ventral wall of the pygofer at about the laterocephalic 
angles of the hypandrium. and they extend directly caudad. They 
are fastened to the connective only at its extreme lateral tip by a 
very narrow band of membrane, although the connection would seem 
to be much greater, since from a ventral view of the organs the con- 
nective covers a large part of the claspers on each side. The union 
of the two can be seen from a ventral and lateral view of the 
genitalia. The styli are fairly well chitinized, but not enough to 
keep them from being flexible. Their function is that of clasping 
or interlocking during copulation. 

The connective (pi. LXI, figs. 7 and 8) is a flat ehitinous plate, 
roughly triangular in shape, which extends between the two claspers 
and is attached to the latter at about the base of their anterior third. 
The lateral margins are produced into a slender recurved hook which 
extends cephalad and wdiose inner margin is attached to the styli. 
The broad basal part of the connective is chitinized, but the apical 
region is entirely membranous. Bordering this membrane on the 
sides are two slender, ehitinous rods which extend caudad, where 
their tapering points are attached to the ventral surface of the base 
of the oedagus. It has been suggested by Doctor Lawson that the 
connective may possibly represent the tenth sternite, since its origin 
has not otherwise been accounted for. 



DoERixc: Lepyronia quadrangularis. 561 

The aedagiis (pi. LXI. figs. 7 and 8) is a term applied to the 
stnioture containing the penis. It has been called by other writers 
the penis sheatii, or merely the penis. It too arises from a genital 
area on the ninth segment. In texture the a3dagus is quite heavily 
chitinized and is very smooth and shiny. Basally the oedagus is 
club-siiaped. flattened dorsoventrally, and gradually narrowing to a 
long, slender, chitinous tube which extends cephalad to a point 
beyond the anterior tips of the styles. Its apex is broadened into a 
flat plate, whose laterocaudal angles bear two longer, slender, taper- 
ing hooks, the penis hooks, which extend directly caudad. 

From the center of the broad plate there appears to arise a 
slender, membranous tube, which also projects caudad, parallel with 
the penis hooks. At the apex of the tube is a circular opening or 
gonoporc. There is also another circular opening at the base of the 
oedagus, which is probably the opening of the ejaculatory duct. 
The flat apical part of the oedagus lies directly beneath the proctiger 
or tenth uromere. The latter has on its lateral margins a small 
hooklike structure (pi. LIX, fig. 2) which seems to fit down around 
the oedagus and which may correspond to the surgonopods of 
Crampton (1922). The oedagus is protective in function, since it 
serves to protect the delicate penis. 

THE FEMALE GENITALIA. 

On the female the genitalia consist of three pairs of appendages, 
which collectively are often spoken of as the ovipositor. More 
properly speaking, however, the ovipositor is made up of only two 
pairs of valves, the ventral and dorsal, and the lateral pair form a 
sheath around them. The eighth sternum of the abdomen, as was 
noted above, is divided into two distinct plates by the processes of 
the ovipositor, while the ninth sternite is represented by two small 
rectangular sclerites to which the lateral valves of the ovipositor are 
attached. 

The lateral valies (pi. LX, fig. 3) have been given other names 
in the literature, such as outer valves, posterior processes, or ovi- 
positor sheaths. They are the outermost of the three pairs and 
fold around the inner pairs as is shown in figure 1. They arise from 
a genital area on the ninth segment and are attached in the adult 
to the vestigial parts of the ninth sternite. The attachment is broad 
and transverse to the caudal end of the sternite. Each lateral 
valve is a broad, spoon-shaped appendage which is deeply concave 
on its inner surface, so that half of it shows from a ventral view 
of the genitalia (pi. LX, fig. 1) and half from a dorsal view (fig. 6). 



562 The University Science Bulletin. 

Its ventral basal part appears to be entirely membranous. The 
pygofers fit very snugly around the lateral valves, but are not at- 
tached to them. These valves are tough, fairly well chitinized and 
with their outer surfaces pubescent. Their ventral surfaces are 
plainly visible from an external view. 

The ventral valves (pi. LX, fig. 2) are the middle pair of valves. 
Other names which have been applied to them are middle valves, 
anterior processes or ventral processes. They arise from the eighth 
sternite, to which they are still attached in the adult. Each valve 
is fastened to its half of the eighth sternite by a membrane which 
connects the mesal corner of the sternite and the cephalomedian 
angle of the valve. They are flat rather broad processes, which 
taper to a fine point and which bear a broad notch near the base of 
the dorsal margin. The ventral valves enfold the dorsal valves and 
the ventral and dorsal valves of each side are fastened together by a 
tongue-and-groove connection. The ventral valves are not as 
heavily chitinized as either the lateral or dorsal ones. 

The dorsal valves (pi. LX, fig. 5), which have also been called 
inner valves or median valves, are the innermost of the three pairs 
of valves, and make up the ovipositor proper. They arise from 
a genital area on the ninth segment of the nymph and are still at- 
tached in the adult stage to the cephalic end of the ninth sternite. 
Each valve is a flat, bladelike structure, broader at the base, but 
tapering to a pointed apex. The basal half of their inner or dorsal 
margins are united and the free apical portions bear teeth. These 
teeth are small, sharply pointed, with broad, shallow indentations 
between them, and are about fifteen in number. The teeth are used 
as a saw with which to cut the plant tissue. The ventral margins of 
these valves bear heavily chitinized grooves into which the tongues 
of the ventral valves are inserted. 

Extending along the approximate median line of the ventral 
valves is a slender, chitinous rodlike structure. Superficially this 
appears to be a chitinous thickening of the valve, but when it is 
traced cephalad it is not found to be attached basally to the sclerite 
to which the valve is fastened. The sclerite to which it is fastened 
is a small, triangular plate (pi. LX, fig. 4), which is attached to the 
lateral margins of the ninth sternite, the cephalic margin of the 
pygofer, and normally lies hidden beneath the eighth sternite. It 
])robably represents the ninth pleurite. 



Doering: Lepyronia quadrangularis. 563 

DEVELOPMENT OF THE CiENITALIA. 

As early as the third iiistar the iiyinphul genitalia arc not only 
distinctly visible, but furnish irliablc characters l)y which to dis- 
tinguish the sexes. They cannot be clearly made out, however, in 
the first two stages without careful study. 

The male genitalia arise from a genital area on the ninth abdom- 
inal segment. In the first and second instars only one pair of 
valves is present. Superficially they appear as two opaque, elongate, 
elevated ridges which occupy the major part of the ninth sternite. 
Closer examination reveals that they are two chitinous pockets 
(pi. LXI, fig. 11, which are separated from each other by a median 
chitinous band and which produce the genital appendages of the 
next nymphal stage. The pockets are attached to the caudal border 
of the genital area with their apices directed caudad and are 
rounded at the tip. The genital area is comparatively short and 
extends cephalad under the caudal margin of the eighth sternum, 
due to the telescopic arrangement of the abdominal segments, so 
that at first glance the pockets appear to arise from the eighth 
sternite. This pair of pockets produces the genital plates of the 
adult. 

In the second instar (fig. 2 I the pockets have increased slightly 
in size, but are similar in other respects to those of the first. 

In the third instar there is a noticeable increase in size of the 
genital area. The pockets of the genital plates have become broader 
but are not so deeply bilobed. In addition to the one pair of pockets 
there is now present another pair, which are located dorsad and 
slightly caudad of the first pair. These produce the genital styles 
or claspers of the adult insect. 

In the fourth instar the genital area has become large and promi- 
nent. There is a great increase in the size of the dorsal pockets, 
which have now become twice as long as the ventral pair. Both 
are rounded at the tip. 

In the fifth instar still greater changes have taken place. The 
ventral pockets have become greatly elongated and their apices have 
diverged slightly. The genital area has lengthened between the 
ventral plates and the dorsal plates so that the former do not ex- 
tend over the latter at all. The dorsal pockets have also diverged 
considerably, and between them can be seen the apices of another 
pair of pockets. These median pockets are only half as long as the 
pockets of the genital styles, are rounded at the apex and produce 
the (I'dagus of the adult. 



564 The University Science Bulletin. 

The jemale genitalia in the first and second instars (pi. LXII, 
figs. 1 and 2) are very difficult to distinguish from those of the male. 
There are two pairs of valves present, but since they are practically 
of the same color and texture and one pair is placed upon the other, 
they appear as a single pair and therefore resemble the male. The 
first pair are small, rounded at the tip and project from the eighth 
sternite as a caudal extension of the latter. Thus it is clearly evi- 
dent that they arise from the eighth sternite, whereas in the male 
the posterior margin of the eighth sternite can be seen to lie on 
top of the first pair of pockets. This pair of pockets develops into 
the ventral valves of the adult ovipositor. The second pair of 
pockets, which produce the dorsal valves of the ovipositor, project 
from above the first pair and are twice as long as the first pair. 
Together they occupy about one-half the length of the ninth 
sternite. 

In the third instar the ventral pockets have increased both in 
width and length. Here they are more easily seen to be attached to 
the eighth sternite. The dorsal pockets have become greatly en- 
larged. They are entirely separated from the ventral pockets, 
occupying about the center of the ninth sternite. The lateral 
pockets are present in the third instar for the first time. They lie 
laterad of the dorsal pockets, are somewhat narrower, although 
much longer, and curve slightly mesad. They develop into the 
lateral valves of the adult. 

In the fourth instar the ventral plates are greatly enlarged, so that 
their apices reach the base of the dorsal valves. The dorsal valves 
are larger than in the preceding instars, but are still the smallest of 
the three pairs. The lateral pockets, which are the largest pair of 
pockets, have their apices projecting much farther caudad than the 
dorsal valves and their bases reaching cephalad almost to the eighth 
sternum. 

In the fifth instar the pockets are very prominent. The ventral 
pockets are broad at the base, extending the entire width of the 
sternum and taper gradually to narrowly rounded apices. The dorsal 
valves are somewhat the shape of the ventral and are entirely covered 
by the latter. Their bases, however, do not reach to the eighth 
sternum. The lateral pockets are fingerlike structures with their 
bases partly covered by the ventral valves and are still slightly 
longer than either of the other two pairs. The genital area is very 
prominent and has been pushed cephalad until it occupies all the 
eighth sternum, and even extends into the seventh, where the indica- 



Doering: Lepyronia quadrangularis. 565 

tion of tlie division of tlie eighth sternum in the adult is shown ])y 
the division of the nyniphal integument. 

From the foi-egoing studies it appears that the male and female 
genitalia are not strictly homologous, since the three pairs of valves 
in the male arise from a genital area on the ninth segment, while in 
the female one pair comes from the eighth and two from the ninth. 
This agrees with Kornhauser's (1919) work on a membracid and 
Hackman's (1923) work on a cicadellid. Kershaw and Muir (1922) 
in their studies of a cercopid, Philcenus leucophthalmus, make the 
statement that the gonopophyses of the male arise in exactly the 
same place as in the female. This conclusion appears to be drawn 
from a study of the fourth and fifth instars only, which are not 
sufficient to give a true interpretation of the origin of these organs. 



566 The University Science Bulletin. 

BIBLIOGRAPHY* 

Amyot. C. J. B., and Serville, J. G. Audixet. Histoire Naturelle ties Insectes. 
Hemipteres, p. LXIII, p. 567; 1843. 

Ashley, K. The Froghopper or Cuckoo Spit on Roses. Review of Apji. Ent., 
vol. VII, ser. A, part 5, p. 209 ; 1919. 

ASHME.AD, W. H. A Proposed Classification of Hemiptera. Ent. Am., 4:67; 
1888. 

B.\KER, Carl F. Notes on Philcenus. Canad. Ent., 29:111-112; 1897. 
Baker, C.\kl F. The Malavan Machaerotinsp (Cercopidae). The Philip])ine 
Joiirn. of Sci., (Sept.) XV". No. 1:67-78; 1919. 

Ball, E. D. A Studv of the Genus Cla,<itoptera. Reprint from la. Acad. 
Science, 3: 182-193; "1896. 

Ball, E. D. A Review of the Cercopidae of North America north of Mexico. 
Rept. la. Acad. Science, pp. 204-226; 1898. 

Ball, E. D. The Food Habits of Some Aphroiihora Larvge. Ohio Nat., 
1:122-124; 1901. 

Ball, E. D. Adaptations to Arid Conditions in Cercopidae and Membracidae. 
Annals Ent. Soc. of America, 8:365-368; 1915. 

Ball, E. D. Notes on Cercoi)id£e. with Descriptions of Some New Species. 
la. Acad, of Sci., 26:143-149; 1919. 

Ball. E. D. Life Cycle in Hemiptera. Annals Ento. Soc. of Am.. 13:143; 1920. 
Barber, George W., and Ellis, Wm. O. Oviposition of Meadow Froghojjper 
and Grass-feeding Froghopper. Psyche, 29, No. 1 ; 1922. 

Boring, Alice M. A Study of Spermatogenesis of Twenty-two Species of 
Membracidae. Jassidae, Cercopidae and Fulgoridae, with special reference to 
the behavior of the Odd Chromosome. J. Exp. Zool. Bait. Md.. 4:469- 
512; 1907. 

Branch, H.-vzel E. The Mori>hologv and Biologv of the Hembracidae of Kan- 
sas, vol. VIII, No. 3:75-111; 1913. 

BuGXioN, E., and Popoff, N. The Mouthparts of the Hemiptera. .\rch. 
Zool. Exp., 47:643-674; 1911. 

CoMSTOcK, John H. and Anna B. Manual for Study of Insects, p. 153; 1895. 

CoMSTOcK, John H., and Kochi, Chijiro. The Skeleton of the Heatl of 
Insects. Amer. Nat., 36:13-45; 1902. 

CRAWFORb, David L. A Monograph of the Jumping Plant Lice or PsvUidae of 

the New World. Smith. Ins. U. S. Nat. Mus. Bull., 85:1-182; 1914. 
Crampton, G. C. Notes on the Thoracic Sclerites of Winged Insects. Ent. 

News., vol. 25, No. 1:15-25; 1914. 
Crampton, G. C. Genitalia and Terminal Aljdominal Structures of Male 

Neuroptera and Mecoptera. with Notes on the Psocidae, Diptera and 

Trichoptera. Psyche, 25. No. 3:47-59; 1918. 

Crampton, G. C. Comparison of Genitalia of Male Hymenoptera, Mecoiitera. 

Neuroptera, Diptera. Trichoptera. Lepidoptera, Homoptera and Streiisij)- 

tera with Those of Lower Insects. Psvche. vol. XXVII, Nos. 2 and 3. 

pp. 34-43. 
Crampton, G. C. The Genitalia of the Males of Certain Hemiptera and 

Homoptera. Bull. Brook. Ento. Soc. vol. XVII, No. 2:46-55; 1922. 
Felt, Ephraim Porter. Twenty-first Report of New York State Entomologist, 

p. 94; 1905. 
Fitch, Asa. Transactions of New York State Agriculture Society, p. 389; 1856. 
Fabre, J. H. The Production of Foam bv Aphrophora. Souvenirs Entomol- 

ogiques. (Septieme Serie.) Paris, 800, pp. 219-233 ; 1900. 

* Only the papers that apply more or less closely to the suhjert matter have l)een used. 



DoERiN(;: Lkpyronia gi adrangularis. 567 

FiNKHorsER, W. 1). Biolotrv of tlie Meiiibiacidir of the Cuvufiii Lake Basin. 
Cornell Univ. Afiiic. Hxp." Sta.. pp. 181-445; 1917. 

Ficc.VKi). W. M. The Systematic Value of tin- Male Genitalia of Delphacidae. 

Ann. Ento. See. Amer., vol. XIV, No. 2, pp. 135-140; 1921. 
GoDiNG, F. W. Synopsis of Subfamilies and Genera of North .\merican 

Cercopida\ Hulf. 111. St. Lab. Nat. Hist.. Ill, p. 483; 1895. 

Garman, Phii.ii'. The Grass-feedinji Frofihojiper or S]iittte Bu>i. (^onn. 
Agric. Kxp. Sta. Bull. 230: 327-334; 1921. 

G.VR.MAN, Phii.m'. Notes on the Life History of Clastoptera obt)i.'<a and 
Lcpyronid quddrattqulnris. .\nn. Knto. Soc. of .Aincr., vol. XVI, No. 2, ])p. 
153-160. 

Gillette, C. P.. and Baker, Carl F. Hemiptera of Colo.. \\. 71; 1895. 

GuFFY, P. L. Report of the Entomologist in charge of Froghopper Investi- 
gation for the Months of October and November, 1914. Rev. of Ai)pl. 
Ento., Ser. A., vol. Ill, part 4:207; 1915. 

Griner, M. Beitrage zur Frage des Aftersecretes der Schaumcicaden. Zool. 
Anz., 23:431-436; 1900. 

H.\CKMAN, Lucy M. Studies on Cicadella hierogly phica . Kan. Univ. Sci. Bull., 
vol. XIV; 1923. 

Harris. T. W. Treatise on Insects Injurious to Vegetation, p. 225; 1862. 

HrxGERFORD. H. B. Biology and Ecology of Aquatic Hemiptera. Univ. of 
Kansas. Sci. Bull., vol. XI. No. 17:1-341; 1919. 

Harper. J. O. Details of Structme of Aphroplioro s]))n)Htria. Sci. Goss.. ])p. 

52-54; 1874. 
Heymoxs, R. Beitrage zur inorphologie und entwicklungsgeschichte der 

Rhynchoten. Nova Acta Acad. Leop. Carol.. Bd. LXXIV, No. 3. Siun- 

inary by author in the Zool. Centrlbl., 7:33-36; 1897. 

Handlirsch, a. Wie viele stigmen haben die Rhynchoten. Ein morpholo- 
gischer beitrag. Verh. ges. Wien.. ])p. 49-510; 1899. Summary, Zool. Cen- 
trabl., VII, ]). 251. 

Hansen, H. J. Gamle og nye kovedmomenter til Cicadariernes morijhologi og 
svstematik. Ent. Tidskr., 11 : 19-76; 1890. Translated bv Kirkaldv in Ento- 
mologist, vol. 33:117, 172, 337; vol. 34:152; vol. 35:216; 1900. 

Kershaw, J. C. Froghoppers. Dept. Agric. Trinidad, Special Circular Nos. 4 
and 5. Summary in Rev. of Appl. Ento., vol. I, ser. A, part 7, p. 233; 1913. 

Kershaw, J. C. The Alimentary Canal of a Cercopid. Psyche, 21:65-71. 

Kershaw, J. C. and Muir, F. The Genitalia of the Auchenorhynchus Homop- 

tera. Annals Ento. Soc. of Amer., vol. XV, No. 3, pp. 201-212; 1922. 
KoRNH.U'SER. Sidney I. The Sexual Characteristics of the Membracid Thclia 

himaculnta (Fabr.). .lourn. of Morph.. vol. 32, No. 3, pp. 531-636; 1919. 

Kirk.\ldy, G. \V. Phylogeny of Homoptera. Canad. Ent.. 42:83; 1910. 

Lawson, Pail B. Cicadellidae of Kansas. Kan. TTniv. Sci. Bull., vol. XII, 
No. 1. ]). 1-373; 1920. 

Lintner, J. A. Ninth Report of New York State Entomologist, p. 393; 1893. 

Meek. Walter J. On the Mouthparts of the Hemiptera. Kan. Univ. Sci 

Bull., vol. II, No. 9:257-277; 1903. 
M.\rl.\tt. C. L. The Hemipterous Mouth. Proc. of Ento. Soc. of Wash., vol. 

3:241-2.50; 1896. 

Metc.\lf, Z. p. The Wing Venation of the Cercopida?. Annals. Ento. Soc. 
Amer.. 10:27-34; 1916. 

Monroe, E. S. A Bubble-blowing Insect. Reprint from Appleton's Popular 
Science Monthly, May, 1900. 

Newell, Anna Gr.ace. The Comparative Morphology of the Genitalia of In- 
sects. Annals of Ent. Soc. Amer., vol. XXI. No. 2:109-136; 1918. 



568 The University Science Bulletin. 

NowELL, W., and Williams, C. B. Sugar-cane Blight in Trinidad: A Summary 
of Conclusions. Bull. Dept. Agric. Trinidad and Tobago, Port of Spain, 
XIX, part 1, pp. 8-10. Summary in Review of Applied Ent., vol. VIII, 
ser. A, part 12, p. 531 ; 1920. 

OsBORN, Herbert. Studies of Life Histories of Froghoppers of Maine. Maine 
Agric. Exp. Sta. Crono. Bull., 254:265-288; 1916. 

PoPENOE. Contribution to a knowledge of the Hemiptera Fauna of Kansas. 
Tr. Kan. Ac, IX:62-64; 1886. 

Readio, p. A. The Ovipositors of Cicadellidse. Kan. Univ. Sci. Bull., vol. 
XIV; 1923. 

RoRER, James Birch. The Green Muscardine of Froghoppers. Reprinted from 
Proc. Agric. Soc. of Trinidad and Tobago, 10:467-482. 

S.-VY, Thomas. The Complete Writings of Thomas Say of Entomology of North 
America. 

Snodgrass, Robert Evans. The Thorax of Insects and the Articulation of the 
Wings. Proc. U. S. Nat. Mus., vol. XXXVI, No. 1687:511-596; 1909. 

Snodgrass, Robert Evans. Seventeen-year Locust. From Smith. Report 
for 1919, pp. 381-409; 1921. 

Smith, John B. The Structure of the Hemipterous Mouth. Science, vol. 19, 
No. 478; 1892. 

Stal, C. Hemip. Fabriciana, IV:54; 1869. 

SuLC, Karel. Respiration, Tracheensystem und Schaumproduktion der 
Schaumcikaden Larvae AphrophorinjB Homoptera. Zs. Wiss. Zool.. 99:147- 

188; 1910. 
Taylor, Leland H. Thoracic Sclerites of Hemiptera. Ann. Ent. Soc. Amer., 

11:225-249; 1918. 
LThler, p. H. Special Descriptive Account of Rhynchota. Rept. U. S. Geol. 

Surv.. 3:355-475 and 765-801; 1877. 
Urich, F. W. Froghoppers in Sugar Cane. Reprint from Bull. Dept. Agric. 

Trinidad, pp. 4-9; 1910. 
Urich. F. W. Sugar-cane Froghopper and Biological Notes on Some Cer- 

coi)ids of Trinidad. Board Agric. Trinidad and Tobago, Circular No. 

9:1-45; 1913. 
UiiicH, F. W. Rearing the Vermilion Froghopper Egg Parasite. Board of 

Agric. Trinidad and Tobago, Circ. No. 7, March, pp. 1-7. Summary Rev. 

of App. Ent., vol. I, ser. A, part 4:116; 1913. 
Uhler, p. R. Checklist of Hemiptera and Heteroptera; 1886. 
Van Duzee, E. P. Hemiptera from Muskoka Lake District of Canada. 

Canad. Ent., 21:8; 1889. 
Van Duzee, E. P. List of Insects Described by Say. Psyche, 5:388; 1890. 
Van Duzee, E. P. Synoptic Table. Trans. Am. Ent. Soc, 19:296; 1892. 
Van Duzee, E. P. Twentieth Rept. N. Y. St. Ent., p. 553; 1905. 
Van Duzee, E. P. Notes on Hemiptera Taken by W. J. Palmer Near Lake 

Temagami, Ont.; 1906. 
Van Duzee, E. P. List of Hemiptera Taken by W. J. Palmer About Quinze 

Lake. Canad. Ent., 40:115; 1908. 
Van Duzee, E. P. Svnonomy of Provancher Collection. Canad. Ent., 44:327; 
1912. 



Doering: Lepyronia quadrangularis. 569 

Van Duzee, E. P. Catalogue of Hemiptora of North America; 1917. 
Walker, E. N. The Terminal Abdominal Structures of Orthopteroid Insects: 

A Phvlogenetic Study. Annals Ent. Soc. Amer., part I, vol. 12, No. 4: 

267-314; 1919. Part II, vol. 15, No. 1:1-67; 1922. 

Williams, C. P. Report of the Froghopper Blight of Sugar Cane in Trinidad, 
No. 1; Jan. 1921. Summarv in Review of App. Ento., vol. IX, ser. A, part 
V, pp. 201-264. 

WiLLL\MS, C. P. Habits' of Tomaspis tristis. Bull. Ent. Res., vol. VII, p. 

271; 1917. 
Williams. C. P. Froghopper Damaging Cacao in Panama. Bull. Ent. Res., 

vol. XIII, part 3, pp. 271-274; 1923. 



)70 The University Science Bulletin. 



PLATE LIV. 

1. Egg. 

2. First instar. 

3. Second instar. 

4. Fifth instar. 

5. Third instar. 

6. Fourth instar. 



PLATE LIV. 




(571) 



PLATE LV. 

1. Lateral view of adult. 

2. Antenna, hishly magnified 

3. Doi-sal A'iew of adult. 

4. Ventral view of adult. 

(572) 



Doerixg: Lepyroxia quadraxgularis. 



573 



PLATE LV 





■ vV^U$"S ~m 




^-^ 






574 The University Science Bulletin. 



PLATE LVL 

1. Dorsal view of head. 

2. Ventral view of head. 

3. Dorsal view of head with vertex removed, showing the tentorium. 

4. Tips of mandible and maxilla, highly magnified. 

5. Lateral view of head. 

6. Lateral view of thorax. 

7. Dorsal view of mesothorax and metathorax. 

8. Ventral view of mesothorax and metathorax. 



Doering: Lepyronia quadrangularis. 



575 



PLATE LVI. 




ironi 

.Vertex 

Ocellui 

Eue 
Olc /puf 
Post (fend 



OcLiptfdl tordrnen 
--hanJibuLv sclerite 

Ldhiu in 





Labrum 

l^anillsrij iclerite / 

Mandibular Stleritc 
LpjpharriK .-- 



Labium 
Sti^leh 

ens 

nfenndl pif 
forearm of tentorium 



posterior arm of 

tentorium 



se of tentorium 



.l^laxilla 



.MmJibU 



.Labrum 

E-pipharunK 



Prescutum 

Scutum 
/ .Scufttlum 




Labium 
'Mandibular Sckrife 
Maxillarii scferife 
5 



Frovotum 



Esimeron 
Epistenium^ 
Trochanfin 
Conn 




Epi mcron, 

Epibferrium, 

M^ron 

Coxa 



Epinieron 

.E pistertiuw 
/ fyesfernur 



Prescufum^ 
icuiu m J 
SiuUllu tn i 
Epimeron . 
Scufu m 3 
■ Stufeliumj 

PoitiLutellum 
Epinierorij 

Sternel lum j 



SfernMm, 



Sierntllum 
Presfcrnu 



Sfcrnurn 




576 The University Science Bulletin. 



PLATE LVII. 

1. Dorsal view of pronotum. 

2. Ventral view of prothorax. 

3. Lateral view of prothorax. 

4. Caudal view of prothorax. 

5. Cephalic view of prothorax. 

6. Lateral view of mesothorax. 

7. Dorsal view of mesothorax. showing wing attachments. 

8. Lateral view of metathorax. 

9. Doi-sal view of metathorax. 

10. Ventral view of mesothorax. 

11. Ventral view of metathorax. 



Doerixg: Lepyroxia quadrangularis. 



577 



PLATE LVII. 




Pronofum 

Sptsfernurn 





rredldre. , 
Anterior phranrn^ 
PrescufuTTi 




Pre vofum 

fp/meroTi.. 

ZpisJcrnuir 

Trochanim j^ , 

r \ 9 rurce 

Coxa 

3 

Preteroife.. 
Pro nolum 

..Fosfcoxale EpnKron... 

-PrecoKale fp'stemuw. 

Trochantin 

...Sfernum , 

Coxd 

Presc iifum.. ^.^y^t^'''^^ ^^ 

\j/^ -/^- ScufuTH 

Pre aid re. \/x /f- -Pas rs /are 

^\\\J fp' rr^ro 71 

Ep.srcrn^rr, \)^ Tltural suture 

Sferimm '^■^:*K^}^^^^--Trccharifm 

,bura/are fi»^r 

T-.-Nofopferale 

i \^\-.. hdanal pttralz 

..Axi I laru cord 
Scufellum 





Wina process 

tp'merov 
Pleural buture 
i-pisternurrt 
Meron 
Coxa. 




Scufurr} 
Fosfscutel/um 



Prcsiernum 
SternuTTj 




Sferne Hum 



578 The University SciencIe Bulletin. 



PLATE LVIII. 

1. Forewing, showing venation. 

2. Mesothoracic leg. 

3. Fore leg. 

4. Fore wing, showing color pattern. 

5. Hind wing. 

6. Dorsal view of tarsal claws. 

7. Hind femur, showing flaplike process. 

8. Metathoracic leg. 

9. Ventral view of tarsal claws. 



Doering: Lepyronia quadrangularis. 



579 



PLATE LVIIl. 




580 The University Science Bulletin. 



PLATE LIX. 

1. Ventral view of male abdomen. 

2. Dorsal view of male abdomen. 

3. Lateral view of male abdomen. 

4. Lateral view of female abdomen. 

5. Dorsal view of female abdomen. 

6. Ventral view of female abdomen. 



DoERIxXG: LePYR0N4A quadrangularis. 



581 



PLATE LIX. 



X sternum 
■jr Sternum 



zpieiirory 




ir pleLtron 






Lateral \/a\ie 
IT Srcrnite. 



xfer^um 
n Pleurov 




n Stcrnife 



I Sfernite 

m Sternite 
I Pleuron 




Ovipositor 



582 The University Science Bulletin. 



PLATE LX. 

1. Ventral view of female genitalia. 

2. Lateral view of ventral valve. 

3. Lateral view of lateral valve. 

4. Rodlike structure found on ventral valve. 

5. Ventral view of dorsal valves and lateral valve. 

6. Dorsal view of female genitalia. 



Doering: Lepyronia quadrangularis. 



583 



PLATE LX. 



fh stcrrtife 




mr fh p'e "r ife. \^- - 




Lateral 
valve 



afh sfernite 
HE pleurite 

' IL pkunte 




aih feraife rv 

lateral valve 

Ventral valve 

Dorsal valvi.. 



)84 The University Science Bulletin. 



PLATE LXI. 

1. Ventral, view of the last three abdominal segments in first-instar male. 

2. Ventral view of last three abdominal segments of second-instar male. 

3. Ventral view of last three abdominal segments of third-instar male. 

4. Ventral view of last three abdominal segments of fourth-instar male. 

5. Lateral view of adult male genitalia. 

6. Ventral view of last three abdominal segments of fifth-instar male. 

7. Ventral view of adult male genitalia. 

8. Dorsal view of adult male genitalia. 



Doering: Lepyroxia quadrangularis. 



585 



PLATE LXI. 




Plate... 
G^r.ifal plafe 

.G^n.ral area. G^mfal stifle 
Geniial orea 



Geniial va/fe 
Genital area 





HH 



fie n I fa/ area 

Genifal stifle- 

\ \ ^Qenifal plat& 




Oedaous 




Genital sfijle 



Connecfive 



O&daaus-.. 

S^i^le 

Plafe... 

Genital area . 





X)eaaQus 



. Gonoporc 

.Cormective 
.Membrane 



jOpeviTio of 
ejaculaforu ducf 



Stale 




586 The University Science Bulletin. 



PLATE LXII. 

1. Ventral view of last three abdominal segments of the first-instar male. 

2. Ventral view of last three abdominal segments of the second-instar 
female. 

3. Ventral view of last three abdominal segments of the third-instar female. 

4. ^"entral view of last three abdominal segments of the fourth-instar 
female. 

5. Ventral view of last three abdominal segments of the fifth-instar female. 

6. Ventral view of the nymphal abdomen with the air channel closed. 

7. Ventral view of the nymphal abdomen with the air channel open. 

8. Caudal view of the tip of the abdomen, showing anal opening. 

9. Mesothoracic leg of a fifth-instar nymph, showing spines on the femur. 

10. First-instar antenna. 

11. Second-instar antenna. 

12. Third-instar antenna. 

13. Fourth-instar antenna. 

14. Fifth-instar antenna. 



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3 2044 093 36' 



863 



Date Due 









1 


L 



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